The objectives of this study were to define the life history of the lodgepole terminal weevil (Pissodes terminalis Hopping) in the Cariboo Forest Region, to quantify the magnitude of damage resulting from weevil attacks, and to determine if P. terminalis activity was associated with measurable tree and stand parameters such as leader dimensions and density. Four study plots in juvenile lodgepole pine (Pinus contorta Dougl. var. latifolia Engelm.) stands near Riske Creek, B. C. were selected to provide data on the biology of the lodgepole terminal weevil. In each plot, weekly collections of 10 weevil-attacked terminal shoots were made from June 3 to August 29, 1980, and monthly collections were taken during September through November. All leaders were dissected and the numbers of each P. terminalis life stage and associated predators/parasitiods were recorded. Seventy-one survey plots in lodgepole pine stands between 8 and 29 years of age throughout the Cariboo Forest Region were assessed for weevil activity. Each survey plot contained 25 trees over 2 metres in height, and each tree was measured for height, diameter at breast height, leader length, and basal leader diameter. Stand age, elevation, density, and all weevil attacks (including year of attack and type of resultant stem deformity) were also recorded. Weevil development appeared to be fairly uniform regardless of stand density and/or elevation; therefore, the data from all four biology study plots were combined to construct a life history diagram. In 46% of the survey plots one or both of the leader dimensions was significantly greater for attacked trees as compared to unattacked trees. In 28% of the plots attacked trees were either significantly taller or larger in diameter. Over 50% of the trees in stands of less than 4,000 stems per hectare had been attacked by P. terminalis at least once. Severe stem deformities such as forks and "stag-heads" comprised a larger proportion of damaged trees in the lower stand density classes. Weevil activity appeared to be higher in 1980 than in 1979. It was concluded that the lodgepole terminal weevil is currently a widely distributed and serious pest of young lodgepole pine stands in the Cariboo Forest Region. Given the present levels of weevil-caused damage as well as the possibility of future attacks, it was recommended that the activity of P. terminalis be considered when silvicultural practices such as juvenile spacing are undertaken in these stands. / Forestry, Faculty of / Graduate

The distribution, population ecology, behavior and host interactions of the root weevil, Hylobius warreni Wood were investigated in lodgepole pine forests in Alberta. Highest incidence of the weevil occurs in the Lower Foothills Section of the Boreal Forest Region, between 2,500 and 4,000 feet in elevation. In even-aged forests weevil numbers are distributed according to stand maturity, stand density, tree size and duff depth. Interactions between these variables modify the patterns of abundance in different stands. Attack incidence upon the host varies, being highest in the collar zone and least on lateral roots. As tree size increases the ratio of weevils on roots/weevils on collar tends to increase. During normal stand development initial invasion of weevils occurs at age 6-10 years, and persists with successive attacks throughout the life of the stand. Weevil populations are highly aggregated in mature stands; "k" values of the negative binomial varied from 0.09 to 0.68, while Taylor's power law gave an aggregation index "b" value of 1.92. Estimates of weevil populations indicated that low levels are characteristic of this species and are maintained, mostly within the range 200-1200 weevils per acre. Estimates of absolute numbers indicate similar levels of abundance occur in young and old stands alike, while population intensity values increase with stand maturity. The highest rate of increase of attack density per tree appears to occur during the ages of 30-45 years. The structure of weevil populations was described and mortality factors were identified and measured for larval, pupal and teneral stages. The main mortality factor of these stages appeared to be from excess moisture in the larval gallery and pupal cell. Only the first 3 larval instars are definable by head capsule width measurement. The feeding behavior of larvae varies with its maturity. In the early instars the feeding pattern relates to bark thickness, but damage is insignificant. Damage of late instars may consist of decorticated gallery lengths up to 24 cm. Larval and pupal habitats are described to indicate the special adaptations for survival. Adults live at least 3 years but lay their eggs during the second and third summers of adulthood. Their seasonal peak of activity occurs in June and early July. Dispersion in the forest tends to be random, commencing about 2 hours after sunset and when temperatures exceed 36-40 °F. Host trees are located partly by vision, the pattern of selection being related to host size. Maximum fecundity per female per season may be 36 or more eggs, but in the field the actual number may not exceed 12.0. Most eggs are deposited singly in niches excavated by the female in the root-collar bark, and are subsequently covered over with excreta. The egg requires a moist environment maintained for up to 42 days for successful hatch. During stand development up to 100 percent of trees may sustain larval feeding damage accumulated to various degrees of intensity. Young trees up to 30 years of age show less resistance to girdling damage than older trees, and reasons are given for this. Estimates of mean height losses of 20-25-year old tree stems sustaining 50 percent girdling were 11.5 and 10.9 percent over 2- and 3-year periods respectively. The total impact of the weevil in the stand as a whole appears to hasten successional changes during stand development. A method of regulation of weevil abundance is postulated and takes into account the behavior of the female during oviposition, host selection, larval feeding habits, cumulative damage and host interactions. Overall numerical restraint and stability of numbers are considered to be effected largely through the inherent behavior of adults. Several weevil control measures are suggested through forest management. Clearcutting of mature timber in alternate strips reduced a weevil population by an estimated 67 percent, but some larvae developed to adults in the cut stumps one and two years after tree removal. The effect of cutting resulted in a concentration of weevils on adjacent trees along stand peripheries, 3-5 years after cutting. / Forestry, Faculty of / Graduate

Dwarf mistletoes are Parasitic flowering plants that infect conifer tree species. The effects of Arceuthobium americanum on the seed crop of Pinus contorta were examined over a two year period to evaluate the effects of altered host reproductive success on host population dynamics. Heavily infected trees produced significantly fewer cones than uninfected trees during the first season in which cones were collected. Cone length, number of seeds per cone, individual seed mass, and total calories per seed were significantly reduced in moderately and heavily infected trees. Trees with different levels of infection showed trends consistent with the above but no significant differences were observed during the second season. The number of seeds that were produced, disseminated, and that germinated per square meter of forest floor was significantly less in heavily infected plots than in lightly infected plots. Survival of one year old seedlings, however, was significantly higher in heavily infected plots. Significant increases in the abundance of woody litter due to increases in mortality of mature trees and witches brooms were positively correlated with increases in seedling survival in heavily infected plots. Survival of trees less than 45 years of age also increased as the level of infection in plots increased. This increases the fraction of total density that saplings represent in heavily infected stands. The total basal area of plots with different levels of infection were uniform, implying that all available environmental resources are being utilized. The differences in age distribution between lightly and heavily infected plots, however, indicate that resources are partitioned differently between saplings and mature trees in plots with different levels of infection. Dominant host trees experienced higher mortality in heavily infected plots than in lightly infected plots. This reduces the total biomass of all mature trees. Increased mortality of dominant host trees frees resources which reduces intraspecific competition thereby contributing to increased sapling survival in heavily infected plots. Consequently, stand density increases as the level of infection increases.

Arceuthobium americanum is a flowering plant which parasitizes Pinus contorta (Lodgepole pine). This study examined branch performance of P. contorta infected to varying degrees with A. americanum.

A study has been conducted with the objectives of (1) identifying weevils and their parasitoids emerging from infested lodgepole pine leaders, (2) determining emergence patterns of hosts and their parasitoids, and (3) obtaining further information on the biologies of the terminal weevils and their natural enemies in British Columbia. The major experiments and biological observations were carried out in young spaced lodgepole pine, (Pinus contorta Dougl. var. latifolia Engelm.), stands at Ellis creek, near Penticton, B.C. A total of 1046 infested leaders were collected. One-third of the terminals were dissected and the numbers of weevils and parasitoids at developmental stages were recorded. The remainder of the leaders were set up for individual rearing. Observations were also made on the feeding and ovipositional behavior of the weevils. Feeding habits of the parasitoids were also studied. Dissections showed that a few adult weevils emerge in the fall of the year of attack. The majority of adults overwinter as larvae but pupation also may occur prior to winter. In addition, dissections indicated that parasitism plays an important role in larval mortality of weevils. Weevils which emerged in the laboratory were identified as being of the following species: Pissodes terminalis Hopping, Magdalis gentilis LeC. and Cylindrocopturus sp. (COLEOPTERA: Curculionidae). M. gentilis is the first weevil species to emerge, in late May. This emergence is followed by that of P. terminalis from early June through mid-July, while Cylindrocopturus sp. emerges from early June through mid-July. P. terminalis attacks the current year's leaders, whereas adult M. gentilis and Cylindrocopturus sp. feed on foliage. All three weevil species utilize lodgepole pine terminal shoots for breeding. Larval feeding under the bark almost always results in the death of the terminal. The terminal weevils have a complex of natural enemies in British Columbia. Parasitoids belong to six families of the order Hymenoptera. The pteromalid Rhopalichus pulchripennis Crawford is the most widely distributed parasitoid species in the province. Two species of Eurytoma (Eurytomidae) ranked second in abundance. Emergence patterns of adult parasitoids are closely synchronized with that of their hosts. Parasitoids were observed feeding on pollen of flowering weeds in the field. This observation suggests that natural parasitoid populations could be enhanced by cultivating lupin, Lupinus sp., in lodgepole pine stands. It was concluded that every effort should be made to minimize weevil numbers in order to prevent formation of crooks, forks and stag-heads. Early emergence of M. gentilis suggests that leader clipping projects should be carried out by early spring. Further research is recommended to ensure correct association between parasitoids and host weevil species and to develop or establish methods for preservation of parasitoids for clipped leaders for release in the forest. / Forestry, Faculty of / Graduate

To prevent the buildup of epidemic level mountain pine beetle populations, conditions of the stand environment they inhabit must be altered. Silvicultural treatment is the most effective means for doing this. Preventative treatments work best when applied while mountain pine beetle populations are still at the endemic population level. Therefore, information necessary for making decisions concerning mountain pine beetle populations in lodgepole pine stands needs to be included in the initial silvicultural prescription planning process, at a time before beetle populations reach outbreak numbers. In this dissertation, several quantitative descriptions of the mountain pine beetle/lodgepole pine relationship were investigated. Models were developed to 1) describe the temperature-dependent development of six mountain pine beetle life stages and 2) describe the amount of loss a stand could sustain if an epidemic level population were to occur in the stand. Concepts of mountain pine beetle risk rating were also discussed. These models and additional information pertaining to the mountain pine beetle/lodgepole pine relationship were incorporated into a knowledge-based system, the MPB Advisory System. This system was designed to help U.S. Forest Service silviculturists include decisions concerning mountain pine beetle populations in the stand management process. / Ph. D.

Arceuthobium americanum is a vascular plant which is parasitic upon Pinus contorta var murrayana. Its documented effects include reductions in host growth, vigor and wood quality. The specific physiological changes that occur in the host are, for the most part, unknown.

Twenty sample plots located in Inonotus tomentosus (Fr.) Teng centers in mixed lodgepole pine (Pinus contorta Dougl. var. latifolia Engelm.) and interior spruce (Picea glauca x engelmannii Engelm.) stands in central British Columbia were studied to investigate the behaviour of, and variation in host species attacked by the pathogen. The effect of infection on the radial increment of spruce and pine was studied by measuring increment bores. Characteristics of the fungus in culture were studied using isolates obtained from each of the study plots. In some cases, the fungus was found to behave differently on spruce and pine. In pine, advanced necrosis of the cambium and phloem of the roots caused crown thinning and eventual standing mortality. In spruce, decay of the heartwood and inner xylem of the roots caused windfall of the tree, often before expression of noticeable crown symptoms. In both host species studied, spread was commonly observed from infected to healthy, contacting roots. Possible cases of spore inoculation through root wounds were also found. In all plots, pine showed a higher infection level than spruce. Subalpine fir (Abies lasiocarpa (Hook.) Nutt.) and Douglas-fir (Pseudotsuqa menziesii (Mirb.) Franco) present within the plots were not found to be infected. Infection by I. tomentosus was not found to reduce significantly the basal area increment in pine or spruce. The fungus in culture demonstrated a large amount of variation in growth rate, mycelium color and mat color and texture. Such variation occurred both within and among isolates, and could not be related to the host species from which the isolate was obtained. / Forestry, Faculty of / Graduate