Defoliating insect systems, defined for the purposes of this thesis as being composed of insects which defoliate forest trees and the species with which they interact, such as their host trees and their natural enemy complexes, exhibit a wide variety of population behaviors. Similarly, a number of theories and models have been proposed to explain these behaviors. These theories emphasize the importance of different ecological processes, often concentrate on the defoliator and overlook the importance of other components. Also, much of the current understanding of the dynamics of these systems has come from forest pest research and management programs, tailored towards specific pest problems and often very short term in nature. This thesis develops and begins to test a general approach for the local dynamics of defoliating insect systems. This framework outlines the system components that are necessary to predict the behavior of defoliating insect systems. It includes ways in which the equilibrium structure of defoliating insect systems, defined as the number of equilibria for each system component, the population levels at which the equilibria occur, and the processes creating the equilibria, might be found. The framework also includes methods of inducing the qualitative behavior of these sys- terns, defined as the periodicity of defoliator outbreaks, the length of outbreaks, and the dynamics of other important system components between, during, and in the decline of defoliator outbreaks. The study begins with a detailed literature review of historical theories of defoliating insect system behavior and of the documented behavior patterns of these systems. Major classes of behavior are identified, as well as the various ecological processes which have been invoked to explain these behaviors. An analysis and documentation of the equilibrium structure and behavior of three defoliating defoliating insect systems, the eastern blackheaded budworm, the eastern spruce budworm, and the jack pine sawfly, are then used to develop general rules about how equilibrium structure and behavior can be explained. This analysis, coupled with the literature review, is used to develop the framework. The framework is then tested against historical defoliator population data and general syntheses of defoliating insect system research to assess its utility and predictability. The major results of the thesis are as follows. First, it appears that the structure and behavior of a defoliating insect system can be explained with five dynamic variables: the abundances of the defoliator; the foliage; the forest; the parasitoid; and the disease; and the effects of weather acting on the defoliator. Second, there appear to be 4 classes of defoliating insect system behavior. Third, the behavior that a defoliating insect system will exhibit seems to be determined by the magnitude of weather effects on defoliator survival and recruitment, the parasitoid numerical response to changing defoliator densities, the disease numerical response to changing defoliator densities, and the vulnerability of the forest to defoliation. Fourth, there seem to be four equilibrium structures the defoliator can exhibit, and one each for the parasitoid, the foliage, the forest, and the disease. Finally, the framework suggests that defoliating insect system structure and behavior can be induced with a particular, well-defined set of information. The framework is successful when applied to particular defoliating insect systems for explaining their behavior, but less successful in explaining defoliator equilibrium structure for other systems. Opportunities for more thorough testing of the framework exist if the particular types of data outlined above are gathered for defoliating insect systems. This lack of data for testing the framework make it currently difficult to clearly define those systems in which the framework is useful and those systems in which it is not. Experiments to test the framework are described and suggestions for future types of applied research on defoliating insect systems are presented. / Science, Faculty of / Zoology, Department of / Graduate

A review of the current understanding of forest tent caterpillar (Malacosoma disstria Hbn.) population dynamics in Ontario suggests that two climatic factors, the temperature at the time of larval feeding and the minimum temperature through the winter, play important roles in determining outbreaks. Comparing the pattern of defoliation to similarly scaled temperature records over 41 years in Ontario shows no relationship between the year to year dynamics of outbreaks and either the temperature through the larval feeding period or the minimum overwintering temperature. A long-term analysis suggests that outbreaks are less severe in those regions with low overwintering temperatures and a patchy distribution of host. This latter finding, combined with an analysis of the synchrony and spread of defoliation, suggests that adult dispersal may play an important role in shaping the dynamics of outbreaks. / Science, Faculty of / Zoology, Department of / Graduate

Thesis (PhD)--Stellenbosch University, 2014. / ENGLISH ABSTRACT: Very little is known about the diversity of fungi associated with Afromontane forests of the Cape Floristic Region (CFR) of South Africa. The ophiostomatoid fungi include many species, some known as pathogens in the CFR, while others are well-known saprophytes important in wood degradation. This study focused on the biodiversity and ecology of tree-associated ophiostomatoid fungi (Ophiostomatales) in the CFR. In addition to this, mites and subcortical beetles associated with the CFR trees were collected, regardless of whether they were associated with ophiostomatoid fungi or not. A relatively high diversity of ophiostomatoid fungi were collected from native trees, ten of which were newly described here. Three further fungal species, two of which are probably new to science, were also collected from exotic Pinus species growing in these forests. Four Ophiostomatales species (including three newly described species) were associated with subcortical beetles on Rapanea melanophloeos and Olea capensis ssp. macrocarpa. These were Sporothrix pallida, Sporothrix aemuluphilus, Raffaelea scabbardiae and Raffaelea rapaneae, associated with the beetles Lanurgus sp. 1, Ctonoxylon sp. 1, Xyleborinus aemuluphilus and a Platypodinae species. This represents a first study to explore the associations between subcortical beetles and ophiostomatoid fungi on native trees in the CFR. In addition to fungi associated with subcortical beetles, several members of the Ophiostomatales associated with wounds on Rapanea melanophloes trees were also collected. These included Ophiostoma stenoceras, Sporothrix reniformis, S. rapaneae, S. lunateae and S. noisomeae. All but O. stenoceras were new to science, and were formally described here. All of these wound-associated species from R. melanophloeos belong to the Sporothrix schenckii – O. stenoceras complex, except for S. noisomeae that was provisionally placed in the S. lignivora complex. Besides fungal taxa collected from wounds on Rapanea melanophloeos, other fungi were also collected from wounds on other host trees species. Three more previously undescribed ophiostomatoid fungal species were collected from this niche. They included Sporothix capensis collected from O. capensis ssp. macrocarpa, Graphilbum roseus collected from many different, unrelated host trees and Graphium ilexiense (Microascales), isolated from wounds on Ilex mitis. The latter represented the first isolation of an ophiostomatoid fungus from this host tree species. Two possibly new fungal species (Sporothrix sp. 1, Ceratocystiopsis sp. 1) and Ophiostoma ips, associated with three bark beetles (Orthotomicus erosus, Hylurgus ligniperda and Hylastes angustatus), were collected from Pinus. Several fungal species were collected from both native trees and non-native trees. These included Sporothrix fusiforme from Brabejum stellatifolium and Acacia mearnsii, O. quercus and O. pluriannulatum-like fungus from several native trees and from A. mearnsii. This suggests a possibility for host shifting of some of these fungi between native and non-native hosts or even between different native hosts. Eight non-ophiostomatoid fungi associated subcortical beetles taxa were found also to infest native trees in the Afromontane forests and in total more than 4500 beetle individuals were collected. Some species of ophiostomatoid fungi collected in this study were found to be associated with other arthropods such as mites. Four phoretic mites species associated with ophiostomatoid fungi (Dendrolaelaps quadrisetus, Histiogaster sp. 3, Elattoma sp. 1 & 2) were collected. In addition, sixteen species of tree wound-associated mites were collected from 12 native trees. Of these, nine were associated with several ophiostomatoid fungi (Graphilbum roseus, O. pluriannulatum-like, O. quercus) that were isolated from several different host trees. This suggests that they may aid in the transport of these fungi from one host species to another. The possible consequences of transfers of Ophiostomatales species between hosts were tested using pathogenicity tests, which highlighted that some fungi are pathogenic on several different trees. Transfers seemed most likely in fungal species isolated from wounds, especially those associated with mites, because the mites may aid in the vectoring of these. When phoretic mites were tested for their specificity to their vector beetles, they proved to be highly specific. Although some of the fungi associated with these mites and their sub-cortical beetles were also pathogenic, it is less likely for these fungi to be transferred to other host tree species due to the high specificity of their arthropod associates. This study represents one of a few studies that focused on ophiostomatoid fungi, subcortical beetles and mites associated with trees in the Afromontane forests of South Africa. Although we collected a high diversity of Ophiostomatales members, many more still await discovery. It is recommended that future studies focus on the complex inter-organismal interactions in many of the systems uncovered in this study. / AFRIKAANSE OPSOMMING: Baie min is bekend oor die diversiteit van fungi wat met die Afromontane woude van die Kaapse Floristiese Streek (KFS) van Suid Afrika geassosieer is. Die ophiostomatoïde fungi sluit baie spesies in, sommiges bekend as patogene in die KFS, terwyl ander bekende en belangrike saprofiete in houtdegradasie is. Hierdie studie het op die biodiversiteit en ekologie van die boom-geassosieerde ophiostomatoïde fungi (Ophiostomatales) in die KFS gefokus. Daarbenewens is myte en subkortikale kewers wat met die KFS bome geassosieer word ook versamel, ongeag of hulle geassosieerd was met ophiostomatoïde fungi of nie. „n Relatief hoë diversiteit van ophiostomatoïde fungi is van inheemse bome versamel, tien waarvan hier nuut beskryf is. Drie verdere fungi spesies, twee waarvan ook waarskynlik nuut is tot die wetenskap, is ook vanaf Pinus spesies versamel wat in hierdie woude gegroei het. Vier Ophiostomatales spesies (insluitend drie nuut beskryfde spesies) wat met subkortikale kewers op Rapanea melanophloeos en Olea capensis L. ssp. macrocarpa geassosieer is, is ook versamel. Hulle was Sporothrix pallida, Sporothrix aemuluphilus, Raffaelea scabbardiae en Raffaelea rapaneae, geassosieer met die kewers Lanurgus sp. 1, Ctonoxylon sp. 1, Xyleborinus aemuluphilus en „n Platypodinae spesie. Hierdie verteenwoordig die eerste studie wat die assosiasies tussen subkortikale kewers en ophiostomatoïde fungi op inheemse bome in die KFS ondersoek. Addisioneel tot fungi geassosieer met die subkortikale kewers, is verskeie lede van die Ophiostomatales vanaf wonde op Rapanea melanophloes bome versamel. Hulle sluit in Ophiostoma stenoceras, Sporothrix reniformis, S. rapaneae, S. lunateae en S. noisomeae. Almal behalwe O. stenoceras was nuut tot die wetenskap, en is hier formeel beskryf. Al hierdie wond-geassosieerde spesies vanaf R. melanophloeos behoort aan die Sporothrix schenckii – O. stenoceras kompleks, behalwe vir S. noisomeae wat voorlopig in die S. lignivora kompleks geplaas is. Benewens fungi taxa wat van die wonde op Rapanea melanophloes versamel is, is ander fungi ook vanaf die wonde op ander gasheer boom spesies versamel. Drie verdere ophiostomatoïde fungus spesies is in hierdie nis versamel. Hulle sluit in Sporothix capensis wat vanaf O. capensis ssp. macrocarpa versamel is, Graphilbum roseus wat vanaf baie verskillende, onverwante gasheer bome versamel is en Graphium ilexiense (Microascales), wat vanaf wonde op Ilex mitis versamel is. Laasgenoemde verteenwoordig die eerste isolasie van „n ophiostomatoïde fungus vanaf hierdie gasheer boom spesie. Twee moontlik nuwe fungus spesies (Sporothrix sp. 1, Ceratocystiopsis sp. 1) en Ophiostoma ips, geassosieer met drie baskewers (Orthotomicus erosus, Hylurgus ligniperda en Hylastes angustatus) is vanaf Pinus versamel. Verskeie fungi spesies is van beide inheemse en nie-inheemse bome versamel. Hulle het Sporothrix fusiforme vanaf Brabejum stellatifolium en Acacia mearnsii, O. quercus en O. pluriannulatum-like fungus vanaf verskeie inheemse bome en vanaf A. mearnsii ingesluit. Dit suggereer die moontlikheid van gasheer-skuiwing van sommige van hierdie fungi tussen inheemse en uitheemse gashere of selfs tussen verskillende inheemse gashere. Agt nie- ophiostomatoïde geassosieerde subkortikale kewers was ook versamel en in totaal is meer as 4500 kewer indiwidue versamel. Sommige ophiostomatoïde fungus spesies wat in hierdie studie versamel is, was met ander geleedpotiges soos myte geassosieer. Vier foretiese myt spesies wat met ophiostomatoïde fungi geassosieer is (Dendrolaelaps quadrisetus, Histiogaster sp. 3, Elattoma sp. 1 & 2), is versamel. Nege addisioneële myt spesies was met verskeie ophiostomatoïde spesies vanaf verskeie boomspesies geassosieer (Graphilbum roseus, O. pluriannulatum-like, O. quercus). Dit suggereer dat myte die vervoer van hierdie fungi van een gasheer spesie na die ander mag bewerkstellig. Die moontlike gevolge van die oordrag van Ophiostomatales spesies tussen gashere is getoets deur patogeniteitstoetse. Dit het beklemtoon dat sommige fungi patogenies is op verskeie onverwante boomspesies. Oordraag van spesies is mees waarskynlik in fungi spesies wat vanaf wonde geisoleer is, veral dié wat met myte geassosieer is, want die myte mag hierdie fungi help vervoer. Toe foretiese myte getoets is vir hulle spesifisiteit tot hulle vektore, is hulle hoogs spesifiek bevind. Alhoewel sommige fungi wat met hierdie myte en hulle geassosieerde kewers geassosieer word wel patogenies is, is dit minder waarskylik dat hulle na ander gasheer bome sal verskuif as gevolg van die hoë spesifisiteit van hulle geleedpotige assosiate. Hierdie studie verteenwoordig een van net enkele studies gefokus op ophiostomatoïde fungi, subkortikale kewers en myte wat met bome van die Afromontane woude van Suid-Afrika geassosieer is. Alhoewel ons „n hoë diversiteit van Ophiotomatale lede versamel het, wag baie meer fungi spesies waarskynlik nog op ontdekking. Daar word voorgestel dat toekomstige studies fokus op die komplekse inter-organismiese interaksies in baie van die sisteme wat in hierdie studie blootgelê is.

The anthracnose pathogen, Colletotrichum gloeosporioides (Penz.) Penz. & Sacc., is a major cause of disease in the avocado industry, causing significant economic losses, and infects all cultivars. In South Africa, Fuerte and Hass varieties are the most widely grown. Identification of genes differentially expressed in avocado during infection with the fungus represents an important step towards understanding the plant’s defence responses and would assist in designing appropriate intervention strategies. In this study, 454 sequencing and analysis of the transcriptome of infected Fuerte avocado fruits were performed using the Roche 454 GS FLX Titanium platform. cDNA libraries enriched for differentially expressed genes were constructed from unharvested and harvested avocado fruit tissues collected after 1, 4 and 24 h post-infection and after 3, 4, 5 and 7 day post-infection, then sequenced.The expression profiles of the genes expressed were measured by a hierarchical clustering algorithm.Subsequently, quantitative real-time PCR was employed to measure the expression of some candidate resistance genes to anthracnose disease and to validate the sequencing results. The single sequencing run produced 215 781 reads from the transcriptome. A total of 70.6 MB of sequence data was generated and subjected to BLAST searches of which about 1500 genes encoding proteins predicted to function in signal transduction, transcriptional control, metabolism, defence, stress response, transportation processes and some genes with unknown functions were identified. The expression profiles studies showed that many expressed genes were either up or down regulated after infection in avocado fruits when compared to the uninfected sample. Salicylic acid and ethylene were identified to be involved in the signalling networks activated in avocado fruit during C. gloeosporioides infection. This study showed that avocado is able to respond to C. gloeosporioides infection by exhibiting a sophisticated molecular system for pathogen recognition and by activating structural and biochemical defence mechanisms.

The shake and wash technique (samples placed in a jar with alcohol added and shaken to remove the mites) was effective in removing 100% of the predator mites, and adult spruce spider mites from Douglas-fir foliage and 98% of the spruce spider mite nymphs. Eighty-eight percent of the spruce spider mite eggs was removed. This technique was considered efficient for removal of spruce spider mite and its predators from Douglas-fir foliage. Sodium hypochlorite (0.84%) added to the alcohol did not increase the number of spruce spider mites or phytoseiid mites removed from the foliage. The intracanopy distribution of spruce spider mite and its predator mites was studied on Douglas-fir Christmas trees in the Willamette Valley, Oregon. When overwintering spruce spider mite egg densities populations were low (<5 eggs/19 cm of stem), significantly more eggs were found on the current season's growth. No significant differences were found between top and bottom halves of the tree or between compass directions. Differences between current and previous season's growth were not found when egg densities were high (>40 eggs/19 cm of stem), but significant differences were found between levels for current season's growth with more eggs found in the upper portion of the canopy. Quadri-directional differences did not exist with either low or high mite populations. Sampling tip or basal stem-halves with low overwintering egg populations did not bias population estimates. Heavy spring rainfall appeared to reduce mite populations as has been reported elsewhere by washing them off the tree and causing increased mortality. Spruce spider mite disperse to the current season's growth shortly after budbreak. Population density rapidly increased in late May and then abruptly declined in mid-July. / Graduation date: 1991

Rodents cause damage in commercial forests by gnawing at the bark of the trees. It is currently estimated that rodent damage in commercial forestry costs the industry R50 million per annum. The species of rodents which cause the damage are not known, neither is the reason behind this behaviour. Through stomach analysis it has been established that 3 species are involved Otomys irroratus, Rhabdomys pumilio and Mastomys natalensis, however this behaviour is confined to the winter. Chemical analysis of the bark reveals that the percentage concentration of nitrogen varies seasonally. The period of high concentration correlates with periods when the natural food of the rodents is restricted and when bark gnawing is most prevalent. In the past the industry's response to the damage has been to treat the areas with rodenticides. Using standard CMR methods, the two commercially-licensed rodenticides and raptor perches were tested to examine their efficacy as rodent control strategies. It was found that at a lower application of I block of rodenticide every third tree there is little difference in the effectiveness of the rodenticide brands and there is also little reduction in the abundance of the rodents. At a higher application rate of I block per tree, the abundance of rodents is reduced but termination of the treatment results in the rodent numbers quickly recovering, indeed they surpassed their original population numbers within 4 months. Apart from the environmental dangers of applying such concentrations of poison, this is clearly an uneconomic solution. My results indicate that at first planting, the sites should be provisioned with raptor perches at a density of 16 ha⁻². Contrary to accepted policy the perches do not require cross pieces, which add to the expense and offer no advantage in raptor residency time. In areas of very high rodent abundance the provisioning of tree collars provide physical protection to the trees. An additional benefit of the collars is that the collars cause a beneficial microclimate around the tree which enhances its growth rate. When the tree is around 2 years old it should be pruned to a height of I ffi, the slash being left in the inter-row. Results show that trees treated in such a way experience no further attack and the rodents browse on the prunings. As the cost of the perches is reduced and the cost of pruning is non-recurring the recommendations provide an economical and environmentally sympathetic alternative to rodenticide application. / Thesis (M.Sc.)-University of Natal, Pietermaritzburg, 1996.

The limited availability of land to forestry and the ensuing emphasis on intensive silviculture, developed a renewed interest in soil pests in the establishment of plantations. Ten field trials were planted over three seasons to determine the mortality factors influencing the establishment of commercial eucalypt and black wattle plantations in the Natal Midlands, and simultaneously, to investigate the chemical control of the soil pest component. A complex of indigenous soil pests contribute to an average 22,9 % failure of Acacia mearnsii and Eucalyptus grandis seedlings from reaching full establishment. This pest complex, which includes termites, whitegrubs, cutworms, tipulid larvae, wireworms, millipedes and nematodes, was responsible for an average 12,3 % of the failure of the plantings to establish. In the absence of termites, in shallow humic soils, whitegrubs followed by cutworms were the most frequent and economically important pests. Eucalypts are more susceptible than wattle seedlings to whitegrub damage when planted in marginal sites. Seedlings in the summer rainfall region were most susceptible to whitegrub damage from December to April; and to cutworm damage during the first two months after planting. An average of 398 hectares was annually damaged by whitegrubs and cutworms. The total annual loss in planting costs and the additional costs of blanking over the three year study period were 1,22 and 2,65 million rands respectively. Existing non-chemical control applicable to woodlot forestry is reported. Chemical control as one of the options in the management of whitegrubs and cutworms was evaluated. The controlled release formulations of carbosulfan 10% and chlorpyrifos 10% at 1,0 g active ingredient/tree (a.i./tree), gamma BRC 0,6% dust at 0,06 g a.i./tree and the synthetic pyrethroid deltamethrin 5 % SC at 0,025 g a.i./tree were persistent and effective in controlling whitegrubs, even when applied early in the planting season. Deltamethrin 5 % SC at 0,025 g a.i./tree was also successful in controlling cutworms. / Thesis (M.Sc.)-University of Natal, Pietermaritzburg, 1995.

Predation by thrips was referred to as early as 1882 when Pergande discussed their habits. Later, Plank (1936) found Leptothrips mali (Fitch) to be predaceous on lepidopterous eggs, and Sharp (1938) cited it as a being a predator of the pecan nut caseborer, Acrobasis caryae Grote. Bailey (1939) stated that the six-spotted thrip, Scolothrips sexmaculatus (Pergande), is predaceous on Tetranychus telarius (L.), Tetranychus pacificus McG., Bryobia praetiosa Koch, Paratetranychus citri McG., Paratetranychus pilosus (C & F.), Paratetranychus indicus Hirst, Paratetranychus yothersi (McG.), Paratetranychus ilicis (McG.), and Phyllocoptes oleivorus Aschm. and in 1943 Bailey and Keifer noted that L. mali fed on tomato russet mite, Phyllocoptes desctructor (Keifer). Muma (1955) discussed the predation of S. sexmaculatus, L. mali, and Aleurodothrips fasciapennis (Franklin) on citrus insects and mites in Florida.

Data for this study were collected by the US Forest Service and the Pennsylvania Bureau of Forestry on nearly 600 plots in central Pennsylvania. Tree and stand characteristics recorded between 1978 and 1985 include estimates of percent defoliation on individual trees. Logistic regression using maximum likelihood estimation was employed to model individual-tree mortality of 15 species in central Pennsylvania that had been defoliated by the gypsy moth. Defoliation was estimated to the nearest ten percent for individual trees. Other variables used for prediction included stand basal area and an individual-tree relative basal area index. Success ranged from no fit for three of the species to an R value (a derivation of Akaike's information criterion) of .613 for white oak. The inclusion of defoliation in the models had a varied effect. For four of the species percent defoliation was not significant. For hickory and white oak respectively, percent defoliation raised the R value by .305 and .290 percentage points. As many as five models for each species were developed: one or two models with no defoliation measure in the model and one each for one, two or three consecutive years of defoliation measures. A beta and gamma function were used to model individual· tree basal area growth for the same 15 species. The models were fit using nonlinear least squares. Variables used include the relative basal area index, stand basal area, site index and a defoliation index that incorporated three years of individual-tree, percent defoliation. The beta and gamma functions fit equally well with values of (1 - relative mean square error) ranging from .1967 to .6290. Results for both models are presented for each species. The defoliation index was a significant variable for five of the fifteen species: white, chestnut, red, and black oak and sassafras. / Master of Science

The effects of gypsy moth defoliation on acorn production (the seed source for oak regeneration), and the forest floor regeneration environment were investigated. Parameters measured included: acorn quantity, weight, development, and viability; litterfall; and forest floor depth and biomass. Nineteen plots in north-central Virginia and western Maryland on which trees suffered heavy, moderate, or no defoliation were monitored over a two-year period. / Master of Science