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Earthworms in Vermont Forest Soils: a Study of Nutrient, Carbon, Nitrogen and Native Plant Responses Ryan Melnichuk University of Vermont

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Earthworms in Vermont Forest Soils: a Study of Nutrient, Carbon, Nitrogen and Native Plant Responses Ryan Melnichuk University of Vermont University of Vermont ScholarWorks @ UVM Graduate College Dissertations and Theses Dissertations and Theses 2016 Earthworms In Vermont Forest Soils: A Study Of Nutrient, Carbon, Nitrogen And Native Plant Responses Ryan Melnichuk University of Vermont Follow this and additional works at: https://scholarworks.uvm.edu/graddis Part of the Ecology and Evolutionary Biology Commons, Plant Sciences Commons, and the Soil Science Commons Recommended Citation Melnichuk, Ryan, "Earthworms In Vermont Forest Soils: A Study Of Nutrient, Carbon, Nitrogen And Native Plant Responses" (2016). Graduate College Dissertations and Theses. 573. https://scholarworks.uvm.edu/graddis/573 This Dissertation is brought to you for free and open access by the Dissertations and Theses at ScholarWorks @ UVM. It has been accepted for inclusion in Graduate College Dissertations and Theses by an authorized administrator of ScholarWorks @ UVM. For more information, please contact [email protected]. EARTHWORMS IN VERMONT FOREST SOILS: A STUDY OF NUTRIENT, CARBON, NITROGEN AND NATIVE PLANT RESPONSES A Dissertation Presented by Ryan Dustin Scott Melnichuk to The Faculty of the Graduate College of The University of Vermont In Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy Specializing in Plant and Soil Science May, 2016 Defense Date: October 29, 2015 Dissertation Examination Committee: Josef H. Görres, Ph.D., Advisor Alison K. Brody, Ph.D., Chairperson Deborah A. Neher, Ph.D. Donald S. Ross, Ph.D. Cynthia J. Forehand, Ph.D., Dean of the Graduate College ABSTRACT Anthropogenic activities surrounding horticulture, agriculture and recreation have increased dispersal of invasive earthworms. The introduction of earthworms initiates many physical and chemical alterations in forest soils previously unoccupied by earthworms. Three trials were performed to assess the effects of earthworms on soil- water dynamics, C and N and defensive/storage compound production by a native plant. The first trial was a greenhouse experiment, performed to assess the impact of two ecologically contrasting invasive earthworm species on percolate and evaporative soil water loss. Mesocosms were constructed to simulate a typical forest Entisol commonly penetrated by the species of interest, Lumbricus terrestris and Amynthas agrestis. Earthworms were added in pair and combination to replicate an average population density observed in Vermont. Percolate water was collected and evaporative water loss balance was recorded over a period of 140 days. C and N were quantified in A-horizon bulk and aggregate soil as well as the subsoil. Residual surface leaf litter was also quantified. Results indicated significantly increased evaporative water loss where either worm was present. Conversely, percolate water loss was significantly reduced in presence L. terrestris alone. C and N analysis revealed that only L. terrestris had a significant effect on aggregate soil C. While the abundance of many herbaceous species is reduced at invasive earthworm sites, Arisaema triphyllum anecdotally have greater densities where earthworms are present. It has been hypothesized that the greater density is caused by a trait that allows this plant to store Ca, often observed at increased concentrations in earthworm invaded soils as Ca-oxalate Here, we tested the hypothesis that oxalate increases in A. triphyllum when earthworms are present. As such, we conducted a two- way factorial greenhouse trial to test whether the changes to soil properties made by two invasive earthworm species (Amynthas agrestis and Lumbricus rubellus) or their physical presence (and bioturbation) had an effect on the plant production of oxalate. Upon quantification of variable soluble oxalate in corms after senescence, we found that earthworm presence increased water soluble and total oxalate significantly as well as marginally significantly in the case of HCl soluble oxalate. No significant changes in oxalate concentrations were observed under soil treatments alone. Carbon and nitrogen are found extensively in both terrestrial and atmospheric cycles. A shift in the equilibrium of these elements can suggest a strong interaction between an introduced variable (invasive earthworms in this case) and the abiotic environment. To better understand changes in soil properties with earthworm invasion, a 112-day mesocosm study was undertaken to examine C and N dynamics. Two epi- endogeic invasive earthworm species Lumbricus rubellus and Amynthas agrestis were selected for study. Greenhouse gas production by total mesocosm and soil were monitored. Gas flux measurements on 11 dates indicate both worm species increase CO 2 and N2O emitted from mesocosm system as well as soil. Mesocosm total C and N (mass balance) indicate significantly more N but no change in C where earthworms are present. This indicates a disruption of denitrification by earthworm invasion that results in increased N2O emissions. This research is the first to examine these variables in concert and confirms holistic view is essential when examining natural systems. ACKNOWLEDGEMENTS I would like to thank my advisor, Josef Görres, for providing guidance and support throughout my research. With his subtle and unrestrictive approach, I was able to follow and explore my results in a manner that would have been unpredictable at the outset of my studies. I am indebted to my committee, Alison Brody, Deborah Neher and Donald Ross for broadening my perspective and challenging my breadth of knowledge such that I should become a more insightful scientist. The high bar set in fundamental knowledge, highly evident passion for continued research in their fields and evanescent hints are qualities I will thus forth aspire to achieve. I would also like to acknowledge my graduate student colleagues, especially Bridgett Hilshey (Jamison) for providing a sounding board as well as words of moral support. Megan, Sebas, Rebecca, Eamon, Lily, Vic, and Tharshani, I hope our paths cross in the future and often. I would also like to acknowledge the efforts of two undergraduate research assistants, Maire Lenihan and Holly Smock; both demonstrated exception detail orientated attention to the tasks performed. And Joel. Coming into the program, I thought I was a master of instrumentation. I quickly realized that was an inaccurate assessment of my skills. I am especially grateful for your patience and support, and hope to carry some of the knowledge and artful mastery of instruments you have shared with me. ii DEDICATION This dissertation is dedicated to the memory of Devin Nahirnak (September 15, 1999 - October 13, 2015), a reminder to treat every day as the only precious commodity… iii TABLE OF CONTENTS Page ACKNOWLEDGEMENTS .............................................................................................. ii DEDICATION ................................................................................................................. iii LIST OF TABLES ......................................................................................................... viii LIST OF FIGURES ..........................................................................................................x CHAPTER 1: COMPREHENSIVE LITERATURE REVIEW ...................................... 1 1.1 Preface................................................................................................................. 1 1.2. Introduction............................................................................................................ 6 1.1.1 History .............................................................................................................. 7 1.1.2 General Behavior and Ecology .......................................................................... 8 1.2 Earthworm Interactions with the Environment....................................................... 9 1.2.1 Physical .............................................................................................................. 9 1.2.2 Biogeochemical ............................................................................................... 11 1.2.3 Plant Interactions ............................................................................................. 15 1.2.4 Other Interactions ............................................................................................ 18 1.3 Earthworms in the Northeastern United States..................................................... 19 1.3.1 Extirpation of Natives and Current Residents ................................................. 19 1.3.2 Species of Interest ............................................................................................ 20 1.3.3 Mechanisms of Introduction and Foothold Success ........................................ 21 1.4 Research Rationale ............................................................................................... 25 iv CHAPTER 2: WORMS AND WATER: A LOOK AT THE INFLUENCE OF TWO INVASIVE EARTHWORMS ON SOIL WATER ....................................................... 26 2.1 Abstract ................................................................................................................. 26 2.2 Introduction........................................................................................................... 27 2.3 Methods and Materials ......................................................................................... 29 2.3.1 Soil and earthworm collections ......................................................................
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