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Thesis (MScAgric)--University of Stellenbosch, 2003. / ENGLISH ABSTRACT: Characterizing the genetic structure of a pest population can provide an understanding of the factors influencing its evolution and assist in its ultimate control. The aim of the present study was to characterize the genetic structure of woolly apple aphid Eriosoma lanigerum (Hausmann) populations in the Western Cape Province in South Africa. Since this economically important apple pest has not previously been characterized at molecular level, it was necessary to evaluate methods for determining the genetic structure of E. lanigerum populations. Two different molecular techniques were evaluated viz. random amplification of polymorphic DNA (RAPD) and amplified fragment length polymorphism (AFLP). This study represents the first application of the latter technique to members of the Aphididae. Aphids were sampled from four regions in the Western Cape in South Africa viz. Elgin, Ceres, Vyeboom and Villiersdorp. A spatially nested sampling design was used to establish the distribution of the genetic variance of aphids. A total of 192 individuals from 13 farms were analysed. Ten RAPD primers were chosen for analysis from an initial assay of 25 after fragment reproducibility had been confirmed. For AFLP analysis three different rare-cutting restriction enzymes were evaluated for AFLP analysis, viz. EcoRI, SseI and MluI. The latter yielded the best results in combination with the frequent-cutting enzyme MseI. Twenty-five AFLP selective primer pairs were evaluated, out of which five were chosen for analysis of the total population. Two hundred and fifty AFLP fragments and 47 RAPD fragments were scored for analysis. Both analyses indicated that a low level of genetic variation was apparent in E. lanigerum populations and that no differentiation resulted from geographic isolation. From RAPD analyses it was deduced that all variation could be attributed to differences between individuals. AFLP analysis indicated that, whereas genetic differences in E. lanigerum populations between orchards were negligible, a significant portion of genetic variation could be attributed to differences between farms and individuals within farms. Therefore, AFLP analysis allowed for finer discrimination of the genetic structure of E. lanigerum populations than RAPD analysis and is recommended for studies of other aphid species. The fact that most of the genetic variation present in E. lanigerum populations could be found on small spatial scales indicated that sampling individuals over a wide geographic area was an ineffective way of detecting the genetic diversity present in E. lanigerum populations. The low level of variation in populations is most likely due to the exclusive occurrence of parthenogenetic reproduction, founder effects (including distribution of infested plant material from a limited source) and selective factors such as the use of resistant rootstocks or pesticides. Furthermore, the low level of variation found indicated that the possibility of controlling E. lanigerum in the Western Cape using host plant resistance is favourable. Thus, plant breeders developing resistance to E. lanigerum can expect plant entries to be exposed to most of the genetic diversity present in Western Cape populations, regardless of location. / AFRIKAANSE OPSOMMING: Die bepaling van die genetiese struktuur van 'n landboukundige plaagpopulasie kan lei tot begrip van die faktore wat die populasie beïnvloed en kan uiteindelike beheer vergemaklik. Die doel van die huidige studie was om die genetiese struktuur van die appelbloedluis Eriosoma lanigerum (Hausmann) in die Wes-Kaap Provinsie van Suid-Afrika te bepaal. Aangesien hierdie belangrike appelplaag nie van tevore op molekulêre vlak bestudeer is nie, was dit nodig om metodes vir die bepaling van die genetiese struktuur van E. lanigerum populasies te evalueer. Twee molekulêre tegnieke is geëvalueer, nl. lukraak geamplifiseerde polimorfiese ONS (RAPD) en geamplifiseerde fragment-lengte polimorfismes (AFLP). Hierdie studie is die eerste om laasgenoemde tegniek te gebruik om lede van die Aphididae te bestudeer. Plantluise is verkry van vier verskillende gebiede in die Wes-Kaap Provinsie van Suid-Afrika nl. Elgin, Ceres, Vyeboom en Villiersdorp. 'n Hierargiese sisteem is gebruik om die verspreiding van die genetiese variasie van plantluise te bepaal. In totaal is 192 individue van 13 plase geanaliseer. Tien RAPD inleiers is gekies uit 'n analise van 25 verskillende inleiers nadat fragment reproduseerbaarheid bevestig is. Drie verskillende restriksie ensieme is geëvalueer vir AFLP analise nl. EcoRI, SseI en Mlul. Die beste resultate is verkry toe MluI saam met MseI gebruik is. Vyf-en-twintig AFLP selektiewe inleier pare is geëvalueer waarvan vyf gekies is vir analise van die totale populasie. Twee-honderd-en-vyftig AFLP fragmente en 47 RAPD fragmente is gedokumenteer vir analise. Beide RAPD en AFLP analises het getoon dat daar 'n lae vlak van genetiese variasie in E. lanigerum populasies is en dat geen differensiasie as gevolg van geografiese isolasie ontstaan het nie. Uit RAPD analise is daar afgelei dat al die variasie toegeskryf kon word aan verskille tussen individue. AFLP het aangetoon dat alhoewel verskille in E. lanigerum populasies tussen boorde laag was, kon 'n hoë persentasie van die variasie toegeskryf word aan verskille tussen plase en individue binne plase. AFLP analise het meer insig in die genetiese struktuur van E. lanigerum populasies verskaf, en word dus aanbeveel vir studies van ander plantluise. Omdat meeste van die genetiese variasie oor klein geografiese afstande verkry word, is steekproefueming oor groot gebiede 'n ondoeltreffende manier om die genetiese variasie binne 'n monster te meet. Die lae vlak van genetiese variasie is waarskynlik te wyte aan partenogenetiese vermeerdering, stigter gevolge (insluitend verspreiding van geïnfesteerde plantmateriaal vanaf 'n beperkte bron), sowel as selektiewe faktore soos die gebruik van bestande onderstokke en insekdoders. Verder dui die lae vlak van variasie aan dat die moontlikheid vir beheer deur gasheerplantbestandheid goed is in die Wes-Kaap. Planttelers kan verseker wees dat hulle plante blootgestel sal wees aan meeste van die genetiese variasie in die Wes-Kaap appelbloedluis populasies ongeag hulle ligging.

Eastern and Carolina hemlock in the eastern United States are experiencing high mortality due to the invasive non-native hemlock woolly adelgid (HWA). The most promising means of control of HWA is the importation of natural enemies from the native range of HWA for classical biological control. Prior to release, natural enemies must be tested for suitability as a control agent, including the ability to locate the target prey. Coleopteran predators, including Scymnus coniferarum and Laricobius osakensis are under consideration as a means of biological control of HWA. Laricobius nigrinus was released in hemlock forests in 2003. It was recently discovered to hybridize with the native Laricobius rubidus. Behavioral responses of these predators to HWA and host tree foliage were observed using a 4-chambered olfactometer, and genetic analysis was used to differentiate responses of L. nigrinus, L. rubidus, and hybrids. In the olfactometer, insects are allowed to amble about the arena and respond to volatile cues from each treatment. Host foliage with and without HWA was tested, as were various comparisons of eastern versus western foliage, host versus non-host foliage, and foliage containing HWA and a congeneric feeding beetle. Olfactometer bioassays demonstrated that foliage from hosts where prey is commonly found is preferable to foliage where prey is seldom found, and that the presence of HWA-induced volatile cues is the strongest driver of behavior, and trumps the presence of a competitor. There is evidence in the study that supports the reliability-detectability phenomenon common in parasitoid biological control agents. Hybrid individuals were found to behave similarly to released L. nigrinus, although in some cases intermediate behavioral traits were evident, with respect to the parental species. This study and others support the continued need for strict testing of potential biological control agents prior to release, as well as a strong impetus for the inclusion and implementation of genetic analysis as a standard component of agent evaluation.

The thesis studied the biological control of woolly aphid (Eriosoma lanigerum Hausm.) using European earwig (Forficula auricularia L.) in an Australian apple orchard IPM program. Woolly aphid populations were monitored over the 1995/96 and 1996/97 growing seasons, completing a four year study of the pest's status and management under IPM programs at Bathurst in the Central Tablelands of NSW. Woolly aphid infestation in 2 IPM treatments, based on alternative control techniques for codling moth (Cydia pomonella L.), namely mating disruption and fenoxycarb, were compared with a conventional insecticide azinphos-methyl program. Further, the thesis studied the toxicity, to adult European earwigs, of chemicals commonly used in Australian apple orchards. The pesticide alpha-cypermethrin is highly toxic to weevil spp. and is used by apple growers in Western Australia for their control, but it is also toxic to earwigs. This trial investigated the potential of alpha-cypermethrin to disrupt the biological control of woolly aphid. Following a single application as a butt spray, alpha-cypermethrin suppressed the number of earwigs in apple trees for 14 weeks. The single application did not reduce earwig numbers to the extent that the biological control of woolly aphid was lost, however a full season program with applications every 14 to 21 days to all trees, as recommended to control weevils, within an orchard is likely to be very disruptive. The research also studied the control of woolly aphid using insecticide root-soil drenches. Imidacloprid provided excellent control of woolly aphid for 3 seasons. Pirimcarb provided some suppression of woolly aphid during the first season, but not in subsequent seasons. Chlorpyrifos and vamidothion failed to control woolly aphid in any season. / Doctor of Philosophy (PhD)

Thesis (M.S.)--West Virginia University, 2007. / Title from document title page. Document formatted into pages; contains vii, 55 p. : ill., maps. Includes abstract. Includes bibliographical references (p. 47-50).

The cotton fleahopper, Pseudatomoscelis seriatus (Reuter) is a widely distributed insect across the United States. Although, it feeds on several native wild hosts, its agricultural importance lies as an economic pest of cotton in several states in the southern United States. No studies have addressed intraspecific genetic and phenotypic variation of this insect pest at a large geographic scale. I examined genetic variation among cotton fleahopper populations associated with cotton in different geographic locations across the southern United States (Chapter II). Using dominant, neutral, nuclear molecular markers (AFLP, amplified fragment length polymorphism) and mitochondrial DNA sequences, I found that overall genetic differentiation among different geographic populations, collected from cotton in eleven cotton growing states, was low but significant. AFLP revealed the presence of three regional groups representing western (Arizona), central (Texas, Oklahoma, Arkansas, Louisiana, Mississippi and Alabama), and eastern (Florida, Georgia, South Carolina and North Carolina) populations. I examined if there were distinct lineages of cotton fleahoppers associated with three of its host plant species: cotton (Gossypium hirsutum), horsemint (Monarda punctata) and woolly croton (Croton capitatus) in five different locations of Texas by using AFLP markers (Chapter III). I found two distinct host-associated lineages at three locations and local panmixia in the other two locations. I tested if host preference of cotton fleahoppers were affected by geographic variation and prior experience. Conducting choice tests with a Y-tube olfactometer, I found that host preference in cotton fleahoppers for horsemint (one of its native host plants) is conserved and unaffected by individual?s prior experience with cotton (Chapter IV). Finally, I explored the role of host-plant species in morphological differentiation of the cotton fleahopper in two locations that differ in presence of distinct host-associated lineages. Using a geometric-morphometric approach, I detected significant effect of host plant and geography on body morphology and wing shape of cotton fleahopper populations (Chapter V). Length of antenna and rostrum were two important traits associated with morphological divergence of cotton and horsemint associated insect populations. Cotton associated individuals had relatively longer antenna and rostrum compared to individuals associated with horsemint.

An invasive insect, hemlock woolly adelgid (HWA), has initiated widespread hemlock decline and mortality in the Great Smoky Mountains National Park (GSMNP). Riparian hemlock mortality impacts on vegetative and aquatic systems of first-order, headwater streams were evaluated. Reference sites for this study were representative of the best available conditions within the GSMNP, with initial stages of HWA presence. Impacted sites were defined as areas with over 90 percent hemlock mortality. Impacted streams had decreased canopy coverage and increased light availability. Residual red maple, yellow birch, and sweet birch capitalized on the loss of hemlock, with increases in relative basal area and species importance values. Rosebay rhododendron responded with increased density and height at impacted sites, thereby preventing woody regeneration. Since long-term regeneration and post-mortality canopy recruitment are limited, alterations of vegetative composition and structure in the stream riparian zone are expected. A seasonal assessment (Sept. 2009 – March 2010) of aquatic impacts revealed increased diurnal variation in stream temperature and exhibited cooler temperatures during the colder months than reference streams. Impacted streams exhibited greater amounts and larger size classes of large woody debris (LWD). Higher concentrations for several nutrients, including silicon, sodium, potassium, magnesium, chlorine, and copper were detected in impacted streams than detected in reference streams. Impacted streams were characterized by higher pH and increased acid neutralization capacity, while reference streams exhibited nitrate concentrations three times higher than impacted stream concentrations. Reference streams were experiencing the initial stage of HWA-induced defoliations, increasing stream nitrate concentrations, while impacted sites had levels suggesting nitrate concentrations have returned to pre-infestation levels. A seasonal assessment of macroinvertebrates found species diversity, abundance, and taxa richness were not affected by hemlock mortality. Impacted streams had a lower density of Chironomidae and Ephemeroptera when compared to reference streams, while Pleuroceridae snails were virtually eliminated at impacted stream sites. Reference streams were dominated by the collector/filter functional feeding group (FFG), while impacted streams were dominated by the scraper FFG. Hemlock mortality induced by the presence of HWA has resulted in short-term impacts to vegetative and aquatic dynamics in stream riparian areas of the Great Smoky Mountain National Park.

The hemlock-silverbell (Tsuga canadensis-Halesia tetraptera) forest type is known to exist in only two places, the Great Smoky Mountains National Park (GRSM) and the Joyce Kilmer National Memorial Forest. The hemlock component of this forest type is currently threatened by the hemlock woolly adelgid (Adelgis tsugae), an invasive aphid-like insect native to Japan. This current status has given rise to the need to investigate the ecological resources of this rare forest type before the hemlock component dies out. The objectives of this study were to determine the nature of the plant/environment and plant/plant associations within this forest type. Within this forest type hemlock was negatively related to protection, aspect, and slope steepness and silverbell was positively related to aspect and slope steepness. This study also identified some interspecific associations such as the negative relationship in the understory between hemlock and striped maple, and provided evidence that understory stems are exhibiting a growth response to hemlock decline in these stands. The information obtained from this study characterizing the plant/environment interactions and even the structural and species components of this forest type will serve as a baseline of data from which to measure change and will provide insight into the mechanisms of species distribution and perhaps into short term scenarios of forest response to hemlock decline and mortality.

Adelges tsugae Annand is a non-native invasive insect threatening the survival of eastern hemlock (Tsuga canadensis) and Carolina hemlock (T. caroliniana). A. tsugae is established in over half of the total range of eastern hemlock and the entire range of Carolina hemlock. Its continued spread, establishment and associated hemlock mortality make research into biological control of A. tsugae crucial. Field surveys of predators associated with A. tsugae in the Pacific Northwest identified a strong correlation between A. tsugae abundance with Laricobius nigrinus and two species of silver fly, Leucopis argenticollis and Leucopis piniperda. Flies in the genus Leucopis are known specialist predators of adelgids and recent studies have shown a strong synchronization between the lifecycles of Leucopis spp. and A. tsugae. The purpose of this study was to test the potential establishment of Leucopis spp. at the southern and northern extent of A. tsugae infested eastern hemlock in eastern United States. In 2015 and 2016, western Leucopis spp. adults were released at two different densities into enclosed branches of A. tsugae infested T. canadensis in Tennessee and New York. A. tsugae on the branches were counted before putting on the enclosure. Four weeks after set-up, all of the enclosures were collected. The number of Leucopis spp. offspring were counted and then stored in ethanol. The number of Leucopis spp. offspring collected were positively related to adelgid density, but did not differ by the number of adult flies per enclosure. Flies collected from enclosures and from the source colony were identified as L.argenticollis and L. piniperda using DNA barcoding. These results show that Leucopis spp. from the Pacific Northwest feed and develop to the adult stage on A. tsugae in the eastern USA. They are able to tolerate environmental conditions during late spring and early summer at the southern and northern extent of the area invaded by A. tsugae in the eastern USA.

Environmental stressors such as nutrient deficiency and insect infestation can significantly impact tree health. Despite much research on the ecological effect on forests in the northeastern United States due to calcium depletion and hemlock woolly adelgid infestation, little is known regarding the physiological mechanisms altered by these stress factors. I tested the hypothesis that calcium depletion, associated with sugar maple decline, compromises water transport processes as a result of calcium-related reductions in cell growth and stabilization. A survey of forest-grown sugar maples from a long-term replicated calcium-manipulation study showed no significant impact of calcium deficiency on wood density, stem hydraulic conductivity (Ks), or vulnerability to cavitation (VC). In vitro removal of xylem-bound calcium showed no impact on VC or air seeding thresholds (Pt). Results suggest that sugar maple decline is not caused by compromises in xylem function due to calcium deficiency. I also tested the hypothesis that hemlock woolly adelgid (Adelges tsugae Annand) (HWA) infestations impact water transport processes and nutrient partitioning in eastern hemlock trees. HWA infestation resulted in higher Ks due to an increase in average tracheid lumen area associated with the proliferation of false rings. HWA-infested trees exhibited higher rates of net photosynthesis and significant changes in foliar nutrient partitioning. These results are the first to demonstrate increases in Ks and alterations in foliar cation levels in response to HWA infestation. In two additional studies, I investigated methods for evaluating the structure and function of xylem networks. Using sequential sectioning of aerial roots of epiphytic aroids, I directly quantified the topographic relation of vessels in a single organ with measurements of vessel length, diameter, vessel end overlap length, and vessel stelar orientation. In a separate study, I explored the relationship between vessel length and measurements of Pt. In establishing guidelines for estimating whole-stem cavitation with the use of single vessel air injection, I demonstrate that calculations of Pt are influenced by stem length measured and removal of native emboli prior to testing. Improvements in tools to quantify xylem structure and function will enhance our ability to understand the responses of forest trees to environmental stress.