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Reticulitermes Flavipes (Isoptera: Rhinotermitidae) Colonies: Reproductive Lifespans, Caste Ratios, Nesting and Foraging Dynamics, and Genetic Architecture

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Reticulitermes Flavipes (Isoptera: Rhinotermitidae) Colonies: Reproductive Lifespans, Caste Ratios, Nesting and Foraging Dynamics, and Genetic Architecture ABSTRACT Title of Dissertation: RETICULITERMES FLAVIPES (ISOPTERA: RHINOTERMITIDAE) COLONIES: REPRODUCTIVE LIFESPANS, CASTE RATIOS, NESTING AND FORAGING DYNAMICS, AND GENETIC ARCHITECTURE Catherine Everitt Long, Doctor of Philosophy, 2005 Dissertation Directed By: Professor Barbara L. Thorne Department of Entomology The eastern subterranean termite, Reticulitermes flavipes (Kollar) (Isoptera: Rhinotermitidae) is a major decomposer of wood and a significant pest of lumber and paper. Despite its economic and ecological importance, key aspects of its colony dynamics are poorly understood. In 1993, laboratory colonies were initiated with alate pairs. In 2003, the colonies contained an average of 12,600 individuals. Ninety-seven percent of kings and 72% of primary queens survived; 29% of colonies contained replacement female reproductives. In addition to providing unprecedented demographic information regarding colony growth rate and longevity, lifespan of founding reproductives, and the response of colonies to the loss of primary kings and queens, these complete colonies demonstrated foraging and nesting activities of socially intact families. The laboratory colonies foraged in multi-resource networks. Travel between the resource nodes was observed, and after 30 weeks all spatial networks were censused. None of the castes was distributed equally among the three resources. Reproductives, which were found in a satellite node in 71% of colonies, and brood did not share the same node a significant portion of the time, suggesting that the nesting strategy was polydomous rather than monodomous. Mark-recapture data indicate that workers were significantly more likely to be found in the resource where they had been located previously, indicating i) they feed non-randomly among the multiple resources and ii) they feed extensively at one location rather than shuttling regularly between satellite- and central nodes (as in a central-place foraging model). Such site fidelity violates an assumption of the Lincoln Index, leading to significant misestimation of actual colony population totals established by census. Worker genotypes, as determined by microsatellite markers, indicate that despite the absence of obvious physical isolation, genetic differentiation had occurred among the workers in one of the queenless colonies. Inbreeding coefficients generated by the queenless colony genotypes did not differ significantly from the predicted F-statistics for colonies of similar breeding structure, confirming that sampled workers can accurately estimate the breeding regime of field populations. RETICULITERMES FLAVIPES (ISOPTERA: RHINOTERMITIDAE) COLONIES: REPRODUCTIVE LIFESPANS, CASTE RATIOS, NESTING AND FORAGING DYNAMICS, AND GENETIC ARCHITECTURE By Catherine Everitt Long Dissertation submitted to the Faculty of the Graduate School of the University of Maryland, College Park, in partial fulfillment of the requirements for the degree of Doctor of Philosophy 2005 Advisory Committee: Professor Barbara L. Thorne, Chair Professor Galen P. Dively Associate Professor William O. Lamp Associate Professor Paula M. Shrewsbury Professor Raymond R. Weil © Copywrite by Catherine Everitt Long 2005 ACKNOWLEDGMENTS It is with love and gratitude that I thank the many family members, friends, and colleagues who inspired and supported me during graduate school. To my peerless mentor, Dr. Barbara Thorne, for encouraging creative exploration and demonstrating a dynamic balance between family and professional responsibilities. To Dr. Nancy Breisch, for being an enthusiastic and resourceful consultant. To my doctoral committee: Drs. Galen Dively, Bill Lamp, Paula Shrewsbury, and Ray Weil, for their high standards and insightful recommendations. To Drs. Larry Douglass and Andie Huberty for their statistical expertise. To Dr. Edward Vargo for generously sharing microsatellite primers, technical expertise, and laboratory facilities. To the Department of Entomology, the Gahan and LaFage families, and American Pest Management, Inc., for their financial support. To my family, for encouragement and support during this extended education. Most of all I thank my wonderful husband, Jeff, for his love and patience, and my beautiful children, William and Lucy, for providing perspective. ii TABLE of CONTENTS ACKOWLEDGEMENTS ................................................................................................... ii LIST of TABLES.................................................................................................................v LIST of FIGURES ..............................................................................................................vi 1 Introduction................................................................................................................ 1 2 Long-term assessment of reproductive lifespan, caste ratios, and colony size in Reticulitermes flavipes................................................................. 3 2.1 Introduction......................................................................................................... 3 2.1.1 General eusocial research............................................................................... 3 2.1.2 Subterranean termite research........................................................................ 4 2.2 Methods............................................................................................................... 6 2.2.1 Culturing techniques and population census.................................................. 6 2.2.2 Statistical analyses ......................................................................................... 8 2.3 Results................................................................................................................. 9 2.3.1 Summary statistics of 8 and 10 year censuses ............................................... 9 2.3.2 Rates of change over 10 years........................................................................ 9 2.3.3 Differences between parental lineages......................................................... 10 2.3.4 Queenright vs. Queenless colonies .............................................................. 16 2.4 Discussion......................................................................................................... 31 2.4.1 Lifespan of primary reproductives............................................................... 31 2.4.2 Colony and individual growth rates............................................................. 31 2.4.3 Number, size, and sex ratio of neotenics ..................................................... 32 2.4.4 Potential genetic basis for caste proportions................................................ 35 2.4.5 Conclusion................................................................................................... 36 3 The impact of multiple, spatially separated food resources on reproduction, caste distribution, and foraging patterns of Reticulitermes flavipes ........................ 38 3.1 Introduction....................................................................................................... 38 3.1.1 General nesting and foraging strategies....................................................... 38 3.1.2 R. flavipes foraging ...................................................................................... 39 3.1.3 Caste distribution......................................................................................... 41 iii 3.2 Methods............................................................................................................. 44 3.2.1 The termites.................................................................................................. 44 3.2.2 Creation of multi-resource colony networks................................................ 44 3.2.3 Travel among the resources ......................................................................... 45 3.2.4 Population distribution................................................................................. 48 3.2.5 Fast marking technique ................................................................................ 50 3.3 Results............................................................................................................... 53 3.3.1 Travel patterns.............................................................................................. 53 3.3.2 Population distribution................................................................................. 56 3.3.3 Resource fidelity.......................................................................................... 58 3.3.4 Population estimation................................................................................... 60 3.4 Discussion......................................................................................................... 64 3.4.1 Practical implications of resource fidelity................................................... 64 3.4.2 Inter-network travel...................................................................................... 68 3.4.3 Distribution of brood and reproductives ...................................................... 71 3.4.4 Polydomy and foraging strategy.................................................................. 71 3.4.5 Conclusion................................................................................................... 72 4 Inbreeding coefficients and breeding structure within laboratory-reared Reticulitermes flavipes colonies: inferences
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