GENETIC VARIATION, LOCAL ADAPTATION AND POPULATION STRUCTURE IN NORTH AMERICAN RED OAK SPECIES, QUERCUS RUBRA L. AND Q. ELLIPSOIDALIS E. J. HILL

Forest trees, like oaks, rely on high levels of genetic variation to adapt to varying environmental conditions. Thus, genetic variation and its distribution are important for the long-term survival and adaptability of oak populations. Climate change is projected to lead to increased drought and fire events as well as a northward migration of tree species, including oaks. Additionally, decline in oak regeneration has become increasingly concerning since it may lead to decreased gene flow and increased inbreeding levels. This will in turn lead to lowered levels of genetic diversity, negatively affecting the growth and survival of populations. At the same time, populations at the species’ distribution edge, like those in this study, could possess important stores of genetic diversity and adaptive potential, while also being vulnerable to climatic or anthropogenic changes. A survey of the level and distribution of genetic variation and identification of potentially adaptive genes is needed since adaptive genetic variation is essential for their long-term survival. Oaks possess a remarkable characteristic in that they maintain their species identity and specific environmental adaptations despite their propensity to hybridize. Thus, in the face of interspecific gene flow, some areas of the genome remain differentiated due to selection. This characteristic allows the study of local environmental adaptation through genetic variation analyses. Furthermore, using genic markers with known putative functions makes it possible to link those differentiated markers to potential adaptive traits (e.g., flowering time, drought stress tolerance). Demographic processes like gene flow and genetic drift also play an important role in how genes (including adaptive genes) are maintained or spread. These processes are influenced by disturbances, both natural and anthropogenic. An examination of how genetic variation is geographically distributed can display how these genetic processes and geographical disturbances influence genetic variation patterns. For example, the spatial clustering of closely related trees could promote inbreeding with associated negative effects (inbreeding depression), if gene flow is limited. In turn this can have negative consequences for a species’ ability to adapt to changing environmental conditions. In contrast, interspecific hybridization may also allow the transfer of genes between species that increase their adaptive potential in a changing environment. I have studied the ecologically divergent, interfertile red oaks, Quercus rubra and Q. ellipsoidalis, to identify genes with potential roles in adaptation to abiotic stress through traits such as drought tolerance and flowering time, and to assess the level and distribution of genetic variation. I found evidence for moderate gene flow between the two species and low interspecific genetic differences at most genetic markers (Lind and Gailing 2013). However, the screening of genic markers with potential roles in phenology and drought tolerance led to the identification of a CONSTANS-like (COL) gene, a candidate gene for flowering time and growth. This marker, located in the coding region of the gene, was highly differentiated between the two species in multiple geographical areas, despite interspecific gene flow, and may play a role in reproductive isolation and adaptive divergence between the two species (Lind-Riehl et al. 2014). Since climate change could result in a northward migration of trees species like oaks, this gene could be important in maintaining species identity despite increased contact zones between species (e.g., increased gene flow). Finally I examined differences in spatial genetic structure (SGS) and genetic variation between species and populations subjected to different management strategies and natural disturbances. Diverse management activities combined with various natural disturbances as well as species specific life history traits influenced SGS patterns and inbreeding levels (Lind-Riehl and Gailing submitted).

Beech bark disease distribution and resistance in Michigan and fungal endophyte ecology of resistant and susceptible beech (Fagus grandifolia Ehrh.)

Beech bark disease (BBD), a non-native association of the fungal pathogen Neonectria faginata and the beech scale insect Cryptococcus fagisuga, has dramatically affected American beech within North American forests. To monitor the spread and effects of BBD in Michigan, a network of forest health monitoring plots was established in 2001 following the disease discovery in Ludington State Park (Mason County). Forest health canopy condition and basic forestry measurements including basal area were reassessed on beech trees in these plots in 2011 and 2012. The influence of bark-inhabiting fungal endophytes on BBD resistance was investigated by collecting cambium tissue from apparently resistant and susceptible beech. Vigor rating showed significant influences of BBD in sample beech resulting in reduced health and substantiated by significant increases of dead beech basal area over time. C. fagisuga distribution was found to be spatially clustered and widespread in the 22 counties in Michigan's Lower Peninsula which contained monitoring plots. Neonectria has been found in Emmet, Cheboygan and Wexford in the Lower Peninsula which may coincide with additional BBD introduction locations. Surveys for BBD resistance resulted in five apparently resistant beech which were added to a BBD resistance database. The most frequently isolated endophytes from cambium tissue were identified by DNA sequencing primarily as Deuteromycetes and Ascomycetes including Chaetomium globosum, Neohendersonia kickxii and Fusarium flocciferum. N. faginata in antagonism trials showed significant growth reduction when paired with three beech fungal endophytes. The results of the antagonism trial and decay tests indicate that N. faginata may be a relatively poor competitor in vivo with limited ability to degrade cellulose.

Carbon in the Peatlands in the Great Lakes Region

Peatlands cover only ~3% of the global land area, but store ~30% of the worlds' soil carbon. There are many different peat types that store different amounts of carbon. Most inventories of carbon storage in northern peatlands have been conducted in the expansive Sphagnum dominated peatlands. Although, northern white cedar peatlands (NW cedar, Thuja occidentalis L.) are also one of the most common peatland types in the Great Lakes Region, occupying more than 2 million hectares. NW cedar swamps are understudied, due in part to the difficulties in collection methods. General lack of rapid and consistent sampling methods has also contributed in a lack of carbon stock quantification for many peatlands. The main objective of this thesis is to quantify: 1) to evaluate peat sampling methods 2) the amount of C-stored and the rates of long-term carbon accumulation in NW cedar peatlands. We sampled 38 peatlands separated into four categories (black ash, NW cedar swamp, sedge, and Sphagnum) during the summers of 2011/2012 across northern MN and the Upper Peninsula of MI. Basal dates of peat indicate that cedar peatlands were between 1970-7790 years old. Cedar peatlands are generally shallower than Sphagnum peat, but due to their higher bulk density, hold similar amounts of carbon with our sites averaging ~800 MgC ha-1. We estimate that NW cedar peatlands store over 1.7 Gt of carbon in the Great Lakes Region. Each of the six methods evaluated had a different level of accuracy and requires varying levels of effort and resources. The depth only method and intermittent sampling method were the most accurate methods of peatland sampling.

Landscape and forest structure at moose mortality sites on Isle Royale National Park. A LiDAR based assessment

Landscape structure and heterogeneity play a potentially important, but little understood role in predator-prey interactions and behaviourally-mediated habitat selection. For example, habitat complexity may either reduce or enhance the efficiency of a predator's efforts to search, track, capture, kill and consume prey. For prey, structural heterogeneity may affect predator detection, avoidance and defense, escape tactics, and the ability to exploit refuges. This study, investigates whether and how vegetation and topographic structure influence the spatial patterns and distribution of moose (Alces alces) mortality due to predation and malnutrition at the local and landscape levels on Isle Royale National Park. 230 locations where wolves (Canis lupus) killed moose during the winters between 2002 and 2010, and 182 moose starvation death sites for the period 1996-2010, were selected from the extensive Isle Royale Wolf-Moose Project carcass database. A variety of LiDAR-derived metrics were generated and used in an algorithm model (Random Forest) to identify, characterize, and classify three-dimensional variables significant to each of the mortality classes. Furthermore, spatial models to predict and assess the likelihood at the landscape scale of moose mortality were developed. This research found that the patterns of moose mortality by predation and malnutrition across the landscape are non-random, have a high degree of spatial variability, and that both mechanisms operate in contexts of comparable physiographic and vegetation structure. Wolf winter hunting locations on Isle Royale are more likely to be a result of its prey habitat selection, although they seem to prioritize the overall areas with higher moose density in the winter. Furthermore, the findings suggest that the distribution of moose mortality by predation is habitat-specific to moose, and not to wolves. In addition, moose sex, age, and health condition also affect mortality site selection, as revealed by subtle differences between sites in vegetation heights, vegetation density, and topography. Vegetation density in particular appears to differentiate mortality locations for distinct classes of moose. The results also emphasize the significance of fine-scale landscape and habitat features when addressing predator-prey interactions. These finer scale findings would be easily missed if analyses were limited to the broader landscape scale alone.

Restoration of forest wetlands: Case studies in Michigan and Finland

Forested wetlands throughout the world are valuable habitats; especially in relatively species-poor northern regions, they can be considered biological hotspots. Unfortunately, these areas have been degraded and destroyed. In recent years, however, the biological importance of wetlands has been increasingly recognized, resulting in the desire to restore disturbed habitats or create in place of destroyed ones. Restoration work is taking place across the globe in a diversity of wetland types, and research must be conducted to determine successful techniques. As a result, two studies of the effects of wetland restoration and creation were conducted in forested wetlands in northern Michigan and southern Finland. In North America, northern white-cedar wetlands have been declining in area, despite attempts to regenerate them. Improved methods for successfully establishing northern white-cedar are needed; as a result, the target of the first study was to determine if creating microtopography could be beneficial for white-cedar recruitment and growth. In northern Europe, spruce swamp forests have become a threatened ecosystem due to extensive drainage for forestry. As part of the restoration of these habitats, i.e. rewetting through ditch blocking, Sphagnum mosses are considered to be a critical element to re-establish, and an in-depth analysis of how Sphagnum is responding to restoration in spruce swamp forests has not been previously done. As a result, the aim of the second study was to investigate the ecophysiological functioning of Sphagnum and feather mosses across a gradient of pristine, drained, and restored boreal spruce swamp forests.

Assessing land use and land cover change in tropical dry forest or Northern Chinandega, Nicaragus from 1985 to 2011

The objective of this study is to gain a quantitative understanding of land use and land cover change (LULCC) that have occurred in a rural Nicaraguan municipality by analyzing Landsat 5 Thematic Mapper (TM) images. By comparing the potential extent of tropical dry forest (TDF) with Landsat 5 TM images, this study analyzes the loss of this forest type on a local level for the municipality of San Juan de Cinco Pinos (63.5 km2) in the Department of Chinandega. Change detection analysis shows where and how land use has changed from 1985 to the present. From 1985 to 2011, nearly 15% of the TDF in San Juan de Cinco Pinos was converted to other land uses. Of the 1434.2 ha of TDF that was present in 1985, 1223.64 ha remained in 2011. The deforestation is primarily a result of agricultural expansion and fuelwood extraction. If current rates of TDF deforestation continue, the municipality faces the prospect of losing its forest cover within the next few decades.

Understanding Michigan nonindustrial private forest landowners attitudes and knowledge about forest management

Approximately one-fourth of the non-industrial private forestland (NIPF) owners in the state of Michigan, who collectively own approximately 50% of the private forested land, have conducted commercial timber harvest in recent years. Previous studies indicated that NIPFs preferred to manage their forest for a sustained yield of high-quality timber, but were limited to even-aged regeneration treatments or conversion for uneven-aged silviculture due to previous cuttings. Improved knowledge about NIPF’s intentions and forest management behavior could be useful for successful implementation of sustained yield management. This study’s objective was to identify more active NIPF’s attitudes towards timber management, their forest management practices and whether their forest management behavior leads or leads not to q management for sustained yield. Active NIPF’s intentions to harvest timber for biofuels and its suitability with NIPF’s forest management behavior will be discussed. Phone interviews of 30 NIPFs who have experience with commercial timber harvests were conducted between August and October 2011. All interviews were recorded, transcribed, and analyzed for identifying NIPF’s motivations, attitudes, forest management behavior and forestry related knowledge. Interviewees, whether consciously or not, tended to manage their land for a sustained yield and they would be willing to harvest timber for biofuels facility as long as it benefits landowners management goals.

A comparison of ground cover and frequency estimation methods for post-harvest soil monitoring

The amount and type of ground cover is an important characteristic to measure when collecting soil disturbance monitoring data after a timber harvest. Estimates of ground cover and bare soil can be used for tracking changes in invasive species, plant growth and regeneration, woody debris loadings, and the risk of surface water runoff and soil erosion. A new method of assessing ground cover and soil disturbance was recently published by the U.S. Forest Service, the Forest Soil Disturbance Monitoring Protocol (FSDMP). This protocol uses the frequency of cover types in small circular (15cm) plots to compare ground surface in pre- and post-harvest condition. While both frequency and percent cover are common methods of describing vegetation, frequency has rarely been used to measure ground surface cover. In this study, three methods for assessing ground cover percent (step-point, 15cm dia. circular and 1x5m visual plot estimates) were compared to the FSDMP frequency method. Results show that the FSDMP method provides significantly higher estimates of ground surface condition for most soil cover types, except coarse wood. The three cover methods had similar estimates for most cover values. The FSDMP method also produced the highest value when bare soil estimates were used to model erosion risk. In a person-hour analysis, estimating ground cover percent in 15cm dia. plots required the least sampling time, and provided standard errors similar to the other cover estimates even at low sampling intensities (n=18). If ground cover estimates are desired in soil monitoring, then a small plot size (15cm dia. circle), or a step-point method can provide a more accurate estimate in less time than the current FSDMP method.

Assessing visual disturbance conditions on the Custer National Forest

Assessment of soil disturbance on the Custer National Forest was conducted during two summers to determine if the U.S. Forest Service Forest Soil Disturbance Monitoring Protocol (FSDMP) was able to distinguish post-harvest soil conditions in a chronological sequence of sites harvested using different ground-based logging systems. Results from the first year of sampling suggested that the FSDMP point sampling method may not be sensitive enough to measure post-harvest disturbance in stands with low levels of disturbance. Therefore, a revised random transect method was used during the second sampling season to determine the actual extent of soil disturbance in these cutting units. Using combined data collected from both summers I detected statistically significant differences (p < 0.05) in fine fraction bulk density measurements between FSDMP disturbance classes across all sites. Disturbance class 3 (most severe) had the highest reported bulk density, which suggest that the FSDMP visual class estimates are defined adequately allowing for correlations to be made between visual disturbance and actual soil physical characteristics. Forest site productivity can be defined by its ability to retain carbon and convert it to above- and belowground biomass. However, forest management activities that alter basic site characteristics have the potential to alter productivity. Soil compaction is one critical management impact that is important to understand; compaction has been shown to impede the root growth potential of plants, reduce water infiltration rates increasing erosion potential, and alter plant available water and nutrients, depending on soil texture. A new method to assess ground cover, erosion, and other soil disturbances was recently published by the U.S. Forest Service, as the Forest Soil Disturbance Protocol (FSDMP). The FSDMP allows soil scientists to visually assign a disturbance class estimate (0 – none, 1, 2, 3 – severe) from field measures of consistently defined soil disturbance indicators (erosion, fire, rutting, compaction, and platy/massive/puddled structure) in small circular (15 cm) plots to compare soil quality properties pre- and post- harvest condition. Using this protocol we were able to determine that ground-based timber harvesting activities occurring on the Custer National Forest are not reaching the 15% maximum threshold for detrimental soil disturbance outlined by the Region 1 Soil Quality Standards.

Retention of canopy trees as biological legacies for balancing woody biomass production and biodiversity in managed aspen forests of the Great Lakes Region

Green-tree retention under the conceptual framework of ecological forestry has the potential to provide both biomass feedstock for industry and maintain quality wildlife habitat. I examined the effects of retained canopy trees as biological legacies (“legacy trees”) in aspen (Populus spp.) forests on above-ground live woody biomass, understory plant floristic quality, and bird diversity. Additionally, I evaluated habitat quality for a high conservation priority species, the Golden-winged Warbler (Vermivora chrysoptera). I selected 27 aspen-dominated forest stands in northern Wisconsin with nine stands in each of three legacy tree retention treatments (conifer retention, hardwood retention, and clearcuts or no retention) across a chronosequence (4-36 years post-harvest). Conifer retention stands had greater legacy tree and all tree species biomass but lower regenerating tree biomass than clearcuts. Coniferous but not hardwood legacy trees appeared to suppress regenerating tree biomass. I evaluated the floristic quality of the understory plant assemblage by estimating the mean coefficient of conservatism (C). Mean C was lower in young stands than in middle-age or old stands; there was a marginally significant (p=0.058) interaction effect between legacy tree retention treatment and stand age. Late-seral plant species were positively associated with stand age and legacy tree diameter or age revealing an important relationship between legacy tree retention and stand development. Bird species richness was greatest in stands with hardwood retention particularly early in stand development. Six conservation priority bird species were indicators of legacy tree retention or clearcuts. Retention of legacy trees in aspen stands provided higher quality nest habitat for the Golden-winged Warbler than clearcuts based on high pairing success and nesting activity. Retention of hardwoods, particularly northern red oak (Quercus rubra), yielded the most consistent positive effects in this study with the highest bird species richness and the highest quality habitat for the Golden-winged Warbler. This treatment maintained stand biomass comparable to clearcuts and did not suppress regenerating tree biomass. In conclusion, legacy tree retention can enhance even-aged management techniques to produce a win-win scenario for the conservation of declining bird species and late-seral understory plants and for production of woody biomass feedstock from naturally regenerating aspen forests.

Managerial perseptions on corporate social responsbility: A Transatlantic comparison between forest products companies in Europe and North America

Corporate Social Responsibility (CSR) addresses the responsibility of companies for their impacts on society. The concept of strategic CSR is becoming increasingly mainstreamed in the forest industry, but there is, however, little consensus on the definition and implementation of CSR. The objective of this research is to build knowledge on the characteristics of CSR and to provide insights on the emerging trend to increase the credibility and legitimacy of CSR through standardization. The study explores how the sustainability managers of European and North American forest companies perceive CSR and the recently released ISO 26000 guidance standard on social responsibility. The conclusions were drawn from an analysis of two data sets; multivariate survey data based on one subset of 30 European and 13 North American responses, and data obtained through in-depth interviewing of 10 sustainability managers that volunteered for an hour long phone discussion about social responsibility practices at their company. The analysis concluded that there are no major differences in the characteristics of cross-Atlantic CSR. Hence, the results were consistent with previous research that suggests that CSR is a case- and company-specific concept. Regarding the components of CSR, environmental issues and organizational governance were key priorities in both regions. Consumer issues, human rights, and financial issues were among the least addressed categories. The study reveals that there are varying perceptions on the ISO 26000 guidance standard, both positive and negative. Moreover, sustainability managers of European and North American forest companies are still uncertain regarding the applicability of the ISO 26000 guidance standard to the forest industry. This study is among the first to provide a preliminary review of the practical implications of the ISO 26000 standard in the forest sector. The results may be utilized by sustainability managers interested in the best practices on CSR, and also by a variety of forest industrial stakeholders interested in the practical outcomes of the long-lasting CSR debate.

Open top chambers and infrared lamps : a comparison of heating efficacy and CO₂/CH₄ dynamics in a Lake Superior coastal peatland

Experimental warming provides a method to determine how an ecosystem will respond to increased temperatures. Northern peatland ecosystems, sensitive to changing climates, provide an excellent setting for experimental warming. Storing great quantities of carbon, northern peatlands play a critical role in regulating global temperatures. Two of the most common methods of experimental warming include open top chambers (OTCs) and infrared (IR) lamps. These warming systems have been used in many ecosystems throughout the world, yet their efficacy to create a warmer environment is variable and has not been widely studied. To date, there has not been a direct, experimentally controlled comparison of OTCs and IR lamps. As a result, a factorial study was implemented to compare the warming efficacy of OTCs and IR lamps and to examine the resulting carbon dioxide (CO2) and methane (CH4) flux rates in a Lake Superior peatland. IR lamps warmed the ecosystem on average by 1-2 #°C, with the majority of warming occurring during nighttime hours. OTC's did not provide any long-term warming above control plots, which is contrary to similar OTC studies at high latitudes. By investigating diurnal heating patterns and micrometeorological variables, we were able to conclude that OTCs were not achieving strong daytime heating peaks and were often cooler than control plots during nighttime hours. Temperate day-length, cloudy and humid conditions, and latent heat loss were factors that inhibited OTC warming. There were no changes in CO2 flux between warming treatments in lawn plots. Gross ecosystem production was significantly greater in IR lamp-hummock plots, while ecosystem respiration was not affected. CH4 flux was not significantly affected by warming treatment. Minimal daytime heating differences, high ambient temperatures, decay resistant substrate, as well as other factors suppressed significant gas flux responses from warming treatments.

Fuelwood collection and consumption: a case study in Lupeta Tanzania

Fuelwood is a major resource in rural areas. Fuelwood collection and consumption habits were monitored in Lupeta, Tanzania through household interviews and fuelwood collection walks. Social dimensions, economic aspects of fuelwood, and alternative fuel sources were also examined. The study found that for all wealth classes, fuelwood is the primary source of fuel used within the village, with the middle and upper classes occasionally supplementing fuelwood with charcoal. Women collect and consume fuelwood for cooking. The majority of women (69%) prefer to use charcoal because fuelwood collection is labor intensive and time consuming. While the use of charcoal would provide more time for other required household activities, local economic constraints inhibit their adoption. The fuel shift from biomass fuels to transition fuels is happening slowly in Lupeta from fuelwood to charcoal. As fuelwood becomes scarcer, improved methods will need to be adopted or the fuel source will change.

Effects of conifer sawdust, hardwood sawdust, and peat on soil properties and a barefoot conifer seedling development

Organic amendments are commonly used to improve tree nursery soil conditions for increased seedling growth. However, few studies compare organic amendments effects on soil conditions, and fewer compare subsequent effects on seedling growth. The effects of three organic amendments on soil properties and seedling growth were investigated at the USDA Forest Service J.W. Toumey Nursery in Watersmeet, MI. Pine sawdust (red pine, Pinus resinosa), hardwood sawdust (maple, Acer spp. and aspen, Populus spp.), and peat were individually incorporated into a loamy sand nursery soil in August, 2006, and soil properties were sampled periodically for the next 14 months. Jack (Pinus banksiana), red, and white pine (Pinus strobus) were sown into test plots in June, 2007 and sampled for growth responses at the end of the growing season. It is hypothesized; pine sawdust and peat can be used as a satisfactory soil amendment to improve soil conditions and produce high quality seedlings, when compared to hardwood sawdust in bareroot nursery soils. This study has the potential to reduce nursery costs while broadening soil amendment options. The addition of peat and pine sawdust increased soil organic matter above control soil conditions after 14 months. However, hardwood sawdust-amended soils did not differ from control soils after same time period. High N concentrations in peat increased total soil N over the other treatments. Similarly, the addition of peat increased soil matric potential and available water over all other treatments. Seedlings grew tallest with the largest stem diameter, and had the largest biomass in both control soil and soil amended with peat, compared to either sawdust treatment. Seedlings grown in peat-amended soils had higher N concentrations than those grown in soils treated with pine sawdust, though neither was different from seedlings grown in control or hardwood sawdust-amended soils. Overall, peat is a well suited organic soil amendment for the enhancement of soil properties, but no amendments were able to increase one-year seedling growth over control soils.

Management impacts on Carex assiniboinensis and Cardamine concatenata on the Ottawa National Forest

Monitoring of herbaceous plants on the Ottawa National Forest (ONF) is used to understand the impact of forest management on understory composition and site conditions. In their planning, national forests are required to take into account management impacts on diversity and ecosystem health. The effect of management on understory species is dependent on various factors, including the intensity of disturbance and the biology of the plant. In the first study in this report, a population of Carex assiniboinensis, a Michigan state threatened species, was monitored for seven seasons including before logging commenced, in order to determine the sedge’s response to a single-tree selection harvest. Analyses provided insights for management of C. assiniboinensis at the stand level over the short-term. In the second study in this report, the use of the cutleaf toothwort (Cardamine concatenata) as a Management Indicator Species on the ONF was reviewed. Data were analyzed to determine the suitability of using C. concatenata to monitor impacts of forest management on site conditions. The various factors that affect understory species population dynamics illuminated the challenges of using indicator species to monitor site conditions. Insights from the study provide a greater understanding of management impacts on understory species across the Ottawa National Forest.