Subterranean Termites of the Oklahoma Tallgrass Prairie Preserve Cross Timbers

Department of Entomology and Plant Patholog

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

THE ART AND SCIENCE OF WOOD: FROM PYROGRAPHY TO TERMITES AND WOOD DECOMPOSITION

Wood is vital to many natural ecosystems, as it provides energy, nutrients, and habitat for organisms from the micro- to the macro- scale. Wood is also critical to humans for similar reasons, and can be an important medium of art and education. This dissertation addresses three diverse aspects of wood with the contexts of science, art, and education. First, we explored the impact of timber harvest techniques and site preparation on microbial wood decay and subterranean termite responses on a forest-stand scale. The amount of coarse woody debris removed post-harvest, coupled with the location and species of the test wood stakes, significantly affected both termite and microbial-mediated decomposition after two and a half years of exposure. These findings help to better understand the impact of timber harvest practices on carbon cycling and associated modes of decay. We then explored effects of wood species and wood surface preparation on pyrography, the art of woodburning. The species of wood and the surface preparation significantly affected line and shading work in pyrography, with more detailed linework produced on hardwoods (Acer rubrum, Populus tremuloides) than on softwoods (Pinus taeda, Pinus strobus). Lastly, placing wood into an educational context, high school level lesson plans that address several science curriculum state and federal benchmarks were developed, to be taught through the active learning technique of pyrography. A general “Introduction to woodburning” lesson plan is included, followed by lesson plans for cellular respiration, human impacts on the environment, photosynthesis, and the carbon cycle. Lesson plans provide instructors with the resources needed to teach across both science and art curriculums. Each lesson plan includes background material, vocabulary, assignments, instructional videos, and PowerPoint presentations. These three chapters weave together science, art, and education using wood as the common thread

The Prairie Naturalist: The Journal of the Great Plains Natural Science Society Volume 50 No. 2

TABLE OF CONTENTS 43 EDITOR’S NOTE RESEARCH ARTICLES 46 Metabolic Gas Emissions from Prairie Soil Containing Foraging Termites Charles E. Konemann, B. M. Kard, Tom A. Royer, and Mark. E. Payton 59 Comparison of northern flying and red squirrel phylogenies with focus on the insular United States Alyssa M. Kiesow, and Hugh B. Britten NOTES 70 Two-Headed White-Tailed Deer Fetus 72 Red-Bellied Snake (Storeria occipitomacilata) Copulation in South Dakota 74 New Breeding Record and Location for Wilson\u27s Phalarope (Phalaropus tricolor) in the Nebraska Great Plains, USA BOOK REVIEWS 76 Sedges and Rushes of Minnesota: The Complete Guide to Species Identification. Welby R. Smith 78 Behavior of the Golden Eagle: An Illustrated Ethogram. David H. Ellis 80 North American Ducks, Geese & Swans: Identification Guide. Frank S. Todd 82 Canids of the World: Wolves, Wild Dogs, Foxes, Jackals, Coyotes, and Their Relatives. José R. Castell

Studies on the Invasion Biology of Social Insects

Social insects (e.g., ants, termites) are among the most prolific group of invasive organisms worldwide. The rapid expansion of both their ecological (i.e., habitats) and global (i.e., countries/continents) distributions has likely been facilitated by the world’s most successful invader – humankind. Therefore, for my PhD research, I performed several investigations into the invasions of social insects to gain further insight into how their invasions have been and are currently being shaped in the Anthropocene. For my first study, I compiled a comprehensive dataset of termite interceptions at US ports of entry spanning the years 1923 to 2017 to elucidate broad patterns in the spread invasive termites to the US. My main findings included a strong regional bias in both the origin (i.e., country/continent) and destination (i.e., port of entry/US region) of interceptions and convincing evidence that invasive termites utilize bridgeheads (i.e., previously invaded locations) to expand their global range. In my next two studies, I reconstructed the invasion histories of two prominent invasive termites – Coptotermes formosanus (native to East Asia) and Reticulitermes flavipes (native to North America). By leveraging existing sample sets previously collected from a large geographic range (i.e., both native and invasive ranges), robust genetic datasets, and approximate Bayesian computation, I inferred a complex invasion history for both species, with multiple invasions from their respective native ranges occurring in conjunction with bridgehead invasions (i.e., invasions originating from a non-native locality). For my final study, I examined Tapinoma sessile’s (odorous house ant) invasion of the urban environment (i.e., cities) from its native natural environments (e.g., forests) across the US. By integrating genetic, chemical, and behavioral data, I discovered strong differentiation between urban and natural populations of the ant in each locality, suggesting cities may be restricting gene flow between habitats and exerting intense selection pressure. Overall, the findings from each of my studies highlight humankind’s powerful and ever-growing influence on the ecological and global distribution of species

Archaeal diversity and community structure in the compartmented gut of highert termites

Phylogenetically higher termites emit higher amounts of methane than lower termites, but the reason for this phenomenon has not been clear. Our comparative study based on 16S rRNA gene sequencing and qPCR analysis of archaeal communities in the guts of higher termites revealed that unlike the lower termites, which host mainly members of Methanobacteriales in their guts, higher termites host a diverse assemblage of methanogenic euryarchaeota comprising representatives of four major orders: Methanobacteriales, Methanosarcinales, Methanomicrobiales and the recently discovered Methanoplasmatales. 16S rRNA-based diversity of archaea was highest in soil-feeding taxa, where nearly all major archaeal groups were represented. Besides the euryarchaeotal lineages, the gut contained also lineages closely related to ammonia-oxidizing Thaumarchaeota and a deep-branching termite specific group of uncultured archaea loosely affiliated to Crenarchaeota. Archaeal diversity in the fungus-cultivating Macrotermes species, in the grass-feeding Trinervitermes sp., and in the wood-feeding Microcerotermes sp., which show low methane emission rates, was much lower. These results show the high methane emission rates in higher termites is reflected in the high diversity, density and complex community structure of archaea in the termite hindguts. Higher termites host gut-specific archaeal communities different from those of lower termites and from other environments and these communities seem to co-evolve with the host termite probably with shift in feeding behavior. Higher termites harbor archaeal lineages which are specific to their gut environment and are different from communities from lower termites and from other environments. Methanogenic archaea are heterogeneously distributed in the highly compartmented gut. Density and diversity of archaea was highest in posterior gut compartments, which also harbored most of the methanogenic activities. The highly alkaline anterior gut compartments were preferentially colonized by Methanosarcinales. Archaeal community structure differed strongly among gut compartments, with communities in the more methanogenic posterior gut sections being distinct from those of the anterior sections, a phenomenon that is reflected in the different micro-environmental conditions in the compartments. Experimental stimulation of methanogenesis in isolated gut sections of soil-feeding termites revealed significant activities of hydrogenotrophic methanogens that are obligately dependent on methanol and formate. Our results suggest that community structure in the different microhabitats is shaped by exogenous factors, such as pH, oxygen status and the availability of methanogenic substrates. The recently discovered Methanoplasmatales are the seventh order of methanogenic euryarchaeota comprising methylotrophic lineages which colonizes higher termite guts and various other environments, and helps to explain the high methane emission rates in higher termites. The methylotrophic nature of termite-derived lineage demonstrates that substrates other than hydrogen drive methanogenesis in higher termites

The phytosanitation of solid wood packaging materials using wood preservatives

New species introductions have been associated with the movement of people for thousands of years. For instance, horses were introduced into North America by Spanish explorers, while pigs were introduced by the Polynesians into many Pacific islands long before the establishment of permanent European settlements. Both of these species introductions resulted in significant changes in both the biological and cultural composition of these places. However, during the past century, the amount of material transported by people has greatly increased due to the globalization of the economy and breakthroughs in material handling technology. These breakthroughs, such as the steel shipping container and the wooden pallet, allow goods to be moved rapidly in a protected environment. This protected environment prevents goods from being damaged while in transit, but it has also improved the ability of pests to survive transit and resulted in invasive species introductions. The volume of goods being transported has steadily increased during the past 50 years and, thus, the number of species introductions has increased as well, resulting in a number of high profile pest introductions. For instance, the introduction of the Asian long horned beetle into Chicago and New York cost millions of dollars to eradicate and resulted in the loss of hardwood trees that previously lined the streets of these cities. In response to the Asian long-horned beetle and the introduction of the pine wood nematode into Europe, the Food and Agriculture Organization of the United Nations drafted and approved International Standard for Phytosanitary Measures Number 15. This measure recognizes solid wood packaging materials as an invasive species pathway and recommends sanitization through heat treatment or fumigation with methyl bromide. However, heat treatment or fumigation are imperfect mitigation tools. Thus, researchers continue to search for alternative methods. Wood preservatives have long been formulated to prevent insect and fungal attack of wood products exposed to warm humid climates, but the ability of these chemicals to eliminate existing insect and fungal colonies has not been investigated. In this study, a number wood preservative systems were investigated to determine if these chemical formulations can be used for this application and to determine if wood preservatives, in general, are suited for use in phytosanitary applications. A method for detecting the presence of wood boring insects through acoustic emissions was investigated. It was thought that a system of microphones and accelerometers could be used to detect wood boring insect presence and activity within a particular wood sample. These assumptions were based on earlier work conducted with termites. This system was to be used to determine if a sample contained wood boring insects prior to using it in the main study. However, acoustic emissions were not useful for this application, since feeding of the wood boring insects was sporadic, unlike termites which feed constantly. Determining the feasibility of using wood preservatives in phytosanitary applications was addressed in three trials: the ability of preservatives to penetrate insect galleries, the ability of the insect larvae and pupae to penetrate a treated barrier and the ability of established wood boring insect populations to complete their life cycle under field conditions in wood pressure treated with preservatives Preservatives completely penetrated between 80-100% of all insect galleries in western redcedar treated with ammoniacal copper quaternary compound, disodium octaborate tetrahydrate, or imidacloprid. However, barriers containing any of these three chemicals failed to prevent larvae from exiting the treated material, even in instances where the barrier was more than 6mm thick. The wood boring insects were unable to complete their life cycle under field conditions in pressure treated wood, while a large number of new house borer adults emerged from the untreated controls. The wood preservatives investigated act more as insecticides than larvacides. However, vacuum pressure impregnation of solid wood packaging materials with the appropriate chemicals could provide lasting protection against invasive species introductions, allowing for the rapid, yet safe transportation of goods around the world

Development of preservative-treated cross-laminated timber and lignin-reinforced polyurethane-adhesive for glued laminated timber

Interest in the use of mass timber in building and construction is growing worldwide, this is due to the structural integrity and reduced environmental footprint of timber-based structures. Concerns associated with the biological and environmental degradation of mass timber necessitate the development of adequate protection strategies to ensure the durability of these products. Preservative treatment is a proven technique that increases the durability and performance of wood in-service and can also be applied to large-sized timber panels such as cross-laminated timber (CLT). Therefore, this study focused on investigating the feasibility of treating prefabricated 3- and 5-layer CLT panels with Copper-azole type C (CA-C) and micronized copper azole (MCA) preservatives. Further, we studied the effects of panel layup and thickness on the preservative impregnation in CLT. Based on the experimental results, we found adequate preservative penetration and retention in the treated 3- and 5-layer CLT panels, particularly in CA-C treated panels. Also, the lengthwise layup shows better treatment results in both CA-C and MCA-treated panels. In addition to the preservative-treatment of CLT panels, this dissertation covers the development of lignin-reinforced polyurethane adhesive (PUR) for bonding glue-laminated timber (Glulam). Herein, the glulam were fabricated and bonded using lignin-reinforced PUR at different wt% (1, 2, and 3) and tested for shear strength, wood failure and delamination. The lignin-treated PUR samples showed improved adhesion properties via high shear strength and reduced delamination compared to the control specimens. Thus, the lignin-reinforced PUR adhesive shows great potential as a bio-based and environment-friendly wood adhesive for producing glulam used in structural applications