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Leaf-Cutter Ant (Atta Cephalotes) Behavioral Ecology of Folivory in a Mixed-Use Guyanan Lowland Rainforest Elizabeth Brianne Karslake University of Missouri-St

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Leaf-Cutter Ant (Atta Cephalotes) Behavioral Ecology of Folivory in a Mixed-Use Guyanan Lowland Rainforest Elizabeth Brianne Karslake University of Missouri-St University of Missouri, St. Louis IRL @ UMSL Theses Graduate Works 4-24-2015 Leaf-cutter Ant (Atta cephalotes) Behavioral Ecology of Folivory in a Mixed-use Guyanan Lowland Rainforest Elizabeth Brianne Karslake University of Missouri-St. Louis, [email protected] Follow this and additional works at: http://irl.umsl.edu/thesis Recommended Citation Karslake, Elizabeth Brianne, "Leaf-cutter Ant (Atta cephalotes) Behavioral Ecology of Folivory in a Mixed-use Guyanan Lowland Rainforest" (2015). Theses. 34. http://irl.umsl.edu/thesis/34 This Thesis is brought to you for free and open access by the Graduate Works at IRL @ UMSL. It has been accepted for inclusion in Theses by an authorized administrator of IRL @ UMSL. For more information, please contact [email protected]. LEAF-CUTTER ANT (ATTA CEPHALOTES) BEHAVIORAL ECOLOGY OF FOLIVORY IN A MIXED-USE GUYANAN LOWLAND RAINFOREST by Elizabeth B. Karslake B.A., Biology, Kalamazoo College, 2012 A THESIS Submitted to the Graduate School of the UNIVERSITY OF MISSOURI- ST. LOUIS In partial Fulfillment of the Requirements for the Degree MASTER OF SCIENCE in BIOLOGY with an emphasis in Animal Behavior May, 2015 Advisory Committee Godfrey R. Bourne, Ph.D. Chairperson Aimee Dunlap, Ph.D. Committee Member Chris Shaffer, Ph.D. Committee Member Karslake, Elizabeth, 2015, UMSL, p. ii Abstract I studied the foraging ecology of a Neotropical leaf-cutter ant, Atta cephalotes, at CEIBA Biological Center, Guyana to elucidate diet choice and foraging strategy. These ants are serous agricultural pests because workers harvest leaves, flowers, fruits, and other plant organs of both cultivated and native plants. The plant materials are used to feed symbiotic fungi whose mycelia tips are the sole food of A. cephalotes larvae. Leaf-cutters were usually found in human disturbed habitats especially slash-and-burned forests cleared for farms, with their higher percentage of sun-exposure and lower plant stem diameters than second growth and primary forests. When given a choice of cultivated and wild plant leaves offered in a randomized smorgasbord test, leaf-cutters accepted significantly more cultivar leaves. These had lower concentrations of secondary compounds than wild plant leaves. In addition, leaf fragment size and thickness transported by returning foragers were related to the foragers’ body length, such that longer ants carried longer and thicker fragments compared to smaller ants. However, there was no relationship between travel distance to the nest and load size, recruitment and returning forager counts, or preference for cultivated plants as predicted by central place foraging theory. In summary, leaf-cutter ants at CEIBA Biological Center were found in human altered forest habitats, exhibited preferences for cultivated over wild plant organs, and did not conform to predictions of central place theory. Therefore, findings have implications for leaf-cutter ant behavioral ecology and agricultural management. ii Karslake, Elizabeth, 2015, UMSL, p. iii Acknowledgements I am especially thankful to Dr. Godfrey R. Bourne, my advisor and director of CEIBA Biological Center because he has been an invaluable mentor since my high-school research experience with the STARS program. Through undergraduate and graduate research projects Godfrey oversaw much of my growth as a scientist and writer. Without his advice, critiques, and encouragement I could not have finished this thesis in a timely manner. Appreciation is expressed to my committee members, Drs. Aimee Dunlap and Chris Shaffer for providing insightful suggestions and encouragement, and expert editing. Thanks to Dr. Rudolf Winter who allowed me access to his lab’s gas-chromatography and mass spectrometry, and he and Joe Kramer assisted in plant volatile extraction and identification. I express gratitude to the CEIBA staff—Joyce Wade, Caroline and Maureen France, they all helped make my stay at CEIBA comfortable. Carolina France, Milford Mathis, and Orrin Clarke generously permitted me to observe leaf-cutter ants destroying their slash-and-burn gardens. Research permits (073013 BR 022 & 60214 BR 016) were issued by the Guyana Environmental Protection Agency. Godfrey, Joyce, Tevin Cooke, and Scheneza Naraine graciously collected ecological measurements in leaf-cutter ant occupied and unoccupied habitats. Furthermore, Tevin and Oswin Ambrose participated in summer 2013 biodiversity course at CEIBA and were awesome research assistants. Tevin returned and assisted during summer 2014 field season, thus, I’m especially thankful for Tevin’s practical suggestions that increased sample sizes during data collection. Sherry Ashanti Naraine and Abigail Marie France, Junior Team Ant Members shadowed me and assisted with data collection when not at school. Karslake, Elizabeth, 2015, UMSL, p. iv Financial support was provided by the Webster Groves Nature Study Society through a Menke Scholarship and CEIBA subsidized my living expenses and provided some research materials and supplies. I am grateful to my parents, Jim and Christine Karslake, for additional finances, encouragement, and sympathies during my graduate studies. Additionally, my 8 year old brother, Jimmy encouraged me to work harder because he always finished his nightly homework before I finished my thesis. Fellow UMSL Biology graduate students, notably Ryan Russell and Guyanpriya (Priya) Maharaj, provided advice about statistics. Priya, Serene Darwish, Hannah Stowe, Oceana O’Dean, and Leanna Kalicharan provided companionship while I was at CEIBA. Finally, I’m grateful to my friends Kellea Floyd, Kamille La Rosa, Elizabeth Sippel, Orel Sosa, and Sandrine Zilikana for encouragement when times were difficult. I could not have completed this thesis without support from fellow Biologists, family, and friends. iv Karslake, Elizabeth, 2015, UMSL, p. v Dedication I dedicate this thesis to Carroll A. Jensen, my maternal grandfather who told everyone about his granddaughter studying to be a scientist just like him. Unfortunately, he died before he could see me graduate. He always reminded me to “work hard, but don’t forget to have fun!” Karslake, Elizabeth, 2015, UMSL, p. vi Table of Contents Page Abstract……………………………………………………………………………. ii Acknowledgments………………………………………………………………… iii Dedication…………………………………………………………………………. v List of Tables………………………………………………………………………. viii List of Figures……………………………………………………………………… ix Chapter I. Impacts of leaf-cutter ants foraging in mixed-use lowland rainforest sites: a review Introduction…………………………………………………………. 1 Ant-Fungus Symbiosis ……………………………………………….. 2 Foraging Behavior…………………………………………………… 5 Plant Species Preferences…………………………………………… 6 Foraging Experiences and Preferences................................................. 10 Optimality Models …………………………………………………... 12 Central Place Foraging Applied to Attini Ants……………………… 14 Preliminary Unpublished Findings and Project Directions……..……... 19 Literature Cited………………………………………………………. 21 Chapter II. Ecological characterization of occupied and unoccupied habitats by the leaf- cutter ant (Atta cephalotes) at CEIBA Biological Center Abstract………………………………………………………………... 24 Introduction…………………………………………………………… 25 Methods ……………………………………………………………….. 27 Results………………………………………………………………… 33 Discussion……………………………………………………………… 37 Literature Cited………………………………………………………... 39 Chapter III. Leaf-cutter ants, Atta cephalotes, already transporting leaves to their nests at CEIBA do not abandon their fragments for domesticated cultivar leaves Abstract……………………………………………………………….. 42 vi Karslake, Elizabeth, 2015, UMSL, p. vii Introduction…………………………………………………………… 43 Methods……………………………………………………………….. 45 Results…………………………………………………………………. 48 Discussion……………………………………………………………… 54 Literature Cited………………………………………………………… 57 Chapter IV. Leaf-cutter ant (Atta cephalotes) plant defoliation strategies do not quite match central place foraging theory Abstract………………………………………………………………… 60 Introduction…………………………………………………………… 61 Methods……………………………………………………………….. 63 Results…………………………………………………………………. 66 Discussion………………………………………………………………70 Literature Cited………………………………………………………... 72 Karslake, Elizabeth, 2015, UMSL, p. viii List of Tables Table 3.1: Plant non-polar metabolites observed in samples of native and forest plants used in randomized smorgasbord........................................ 53 viii Karslake, Elizabeth, 2015, UMSL, p. ix List of Figures Figure 2.1: A. cephalotes castes...................................................................................30 Figure 2.2: Comparison of slash-and-burn agriculture plots with leaf-cutter ants and old second growth forests lacking ants.....................................................30 Figure 2.3: Number of leaf-cutter ant nest entrances in habitats with and without active nests................................................................................................34 Figure 2.4: Percent sun exposure of sampled sites in habitats with either active or inactive A. cephalotes nests......................................................................35 Figure 2.5: Diameter at breast height of plants in habitats with either active or inactive A. cephalotes nests......................................................................36 Figure 3.1: Median discovery times for leaf treatments presented during randomized smorgasbords.............................................................................................50 Figure 3.2: Median handling times for leaf treatments presented during randomized smorgasbords.............................................................................................51
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