INTEGRATED MANAGEMENT OF THE INVASIVE SPECIES CIRSIUM ARVENSE, CANADA THISTLE, AND PHRAGMITES AUSTRALIS, COMMON REED: USING ECOLOGICALLY BASED CONTROL MEASURES AT EACH STAGE OF THE INVASION PROCESS

Invasive plant species threaten natural areas by reducing biodiversity and altering ecosystem functions. They also impact agriculture by reducing crop and livestock productivity. Millions of dollars are spent on invasive species control each year, and traditionally, herbicides are used to manage invasive species. Herbicides have human and environmental health risks associated with them; therefore, it is essential that land managers and stakeholders attempt to reduce these risks by utilizing the principles of integrated weed management. Integrated weed management is a practice that incorporates a variety of measures and focuses on the ecology of the invasive plant to manage it. Roadways are high risk areas that have high incidence of invasive species. Roadways act as conduits for invasive species spread and are ideal harborages for population growth; therefore, roadways should be a primary target for invasive species control. There are four stages in the invasion process which an invasive species must overcome: transport, establishment, spread, and impact. The aim of this dissertation was to focus on these four stages and examine the mechanisms underlying the progression from one stage to the next, while also developing integrated weed management strategies. The target species were Phragmites australis, common reed, and Cisrium arvense, Canada thistle. The transport and establishment risks of P. australis can be reduced by removing rhizome fragments from soil when roadside maintenance is performed. The establishment and spread of C. arvense can be reduced by planting particular resistant species, e.g. Heterotheca villosa, especially those that can reduce light transmittance to the soil. Finally, the spread and impact of C. arvense can be mitigated on roadsides through the use of the herbicide aminopyralid. The risks associated with herbicide drift produced by application equipment can be reduced by using the Wet-Blade herbicide application system.

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.

Compensatory growth in moose and its relationship to sex, longevity, intraspecific competition, and senescence

Individual life history theory is largely focused on understanding the extent to which various phenotypes of an organism are adaptive and whether they represent life history trade-offs. Compensatory growth (CG) is increasingly appreciated as a phenotype of interest to evolutionary ecologists. CG or catch-up growth involves the ability of an organism to grow at a faster-than-normal rate following periods of under-nutrition once conditions subsequently improve. Here, I examine CG in a population of moose (Alces alces) living on Isle Royale, a remote island in Lake Superior, North America. I gained insights about CG from measurements of skeletal remains of 841 moose born throughout a 52-year period. In particular, I compared the length of the metatarsal bone (ML) with several skull measurements. While ML is an index of growth while the moose is in utero and during the first year or two of life, a moose skull continues to grow until a moose is approximately 5 years of age. Because of these differences, the strength of correlation between ML and skull measurements, for a group of moose (say female moose) is an indication of that group’s capacity for CG. Using this logic, I conducted analyses whose results suggest that the capacity for CG did not differ between sexes, between individuals born during periods of high and low population densities, or between individuals exhibiting signs of senescence and those that do not. The analysis did however suggest that long-lived individuals had a greater capacity for CG than short-lived individuals. These results suggest that CG in moose is an adaptive trait and might not be associated with life history trade-offs.

Predator-prey: interactions between the spiny waterflea (Bythotrephes longimanus) and pumpkinseed sunfish (Lepomis gibbosus)

Small pumpkinseed sunfish ( Lepomis gibbosus), were found to be capable of removing the spine of Bythotrephes longimanus, an invasive cladoceran. Because fish consumption may be important in the dispersal or control of Bythotrephes, aquarium feeding experiments were conducted to 1) establish if the spine removal behavior of the pumpkinseeds was locally unique; 2) quantify how frequently pumpkinseeds exhibit the behavior; 3) determine if pumpkinseed handle Bythotrephes more quickly than other species of fish; and 4) verify if Bythotrephes' resting eggs pass through the digestive systems of pumpkinseeds in viable condition. The experiments revealed that pumpkinseeds (45-70 mm TL) from two geographic regions were more successful (100%) at removing Bythotrephes' spine, and handled Bythotrephes more quickly than yellow perch (Perca flavescens) (49-57 mm TL) and smallmouth bass (Micropterus dolomieu) (50-57mm TL) used in the study. Of 244 live Bythotrephes' resting eggs fed to the pumpkinseeds, 104 (42.6%) passed through their digestive systems. From those eggs, only 10 successfully hatched. Preliminary enclosure experiments were carried out and indicated that pumpkinseeds will consume Bythotrephes in natural settings. These findings provide new evidence that certain fish, with specialized morphology for prey manipulation, have the ability to influence the distribution and establishment of Bythotrephes.

INTERACTIVE CONTROLS OF WATER TABLE POSITION AND PLANT FUNCTIONAL TYPES ON PEAT POREWATER CHARACTER IN NORTHERN BOG ECOSYSTEMS: IMPLICATIONS FOR CARBON CYCLING DYNAMICS

Northern wetlands, and particularly peatlands, have been shown to store around 30% of the world's soil carbon and thus play a significant role in the carbon cycle of our planet. Changes in climate are altering peatland hydrology and vegetation communities. These changes are possibly resulting in declines in the ability of peatlands to sequester carbon because losses through carbon oxidation and mineralization are likely to increase relative to C inputs from net primary production in a warmer, drier climate. However, the consequences of interactive effects of altered hydrology and vegetation on carbon storage are not well understood. This research evaluated the importance of plant species, water table, and their interactive effects on porewater quality in a northern peatland with an average pH of 4.54, ranging from 4.15 to 4.8. We assessed the effects of plant functional group (ericaceous shrubs, sedges, and bryophytes) and water table position on biogeochemical processes. Specifically, we measured dissolved organic carbon (DOC), total dissolved nitrogen (TDN), potential enzyme activity, organic acids, anions and cations, spectral indexes of aromaticity, and phenolic content. Our results indicate that acetate and propionate concentrations in the sedge-dominated communities declined with depth and water table drawdown, relative to the control and ericaceous treatments. DOC increased in the lowered water table treatments in all vegetation community types, and the peat porewater C:N ratio declined in the sedge-dominated treatments when the water table was lowered. The relationship between DOC and ferrous iron showed significant responses to vegetation type; the exclusion of Ericaceae resulted in less ferrous iron per unit DOC compared to mixed species treatments and Ericaceae alone. This observation was corroborated with higher mean oxidation redox potential profiles (integrating 20, 40, and 70 cm) measured in the sedge treatments, compared with the mixed and Ericaceae species treatments over a growing season. Enzymatic activities did not show as strong of a response to treatments as expected; the oxidative enzyme peroxidase and the hydrolytic enzyme phosphatase were the only enzymes to respond to water table, where the potential activity of both enzymes increased with water table drawdown. Overall, there were significant interactive effects between changes in vegetation and water table position on peat porewater composition. These data suggest that vegetation effects on oxidation reduction potentials and peat porewater character can be as important as water table position in northern bog ecosystems.

INVESTIGATING THE IMPACTS OF STREAMBED HABITAT HETEROGENEITY ON ECOSYSTEM STRUCTURE AND PROCESSES USING BASIC AND APPLIED PERSPECTIVES

Stream restoration often focuses on increasing habitat heterogeneity to reverse ecosystem degradation. However, the connection between heterogeneity and ecosystem structure and processes is poorly understood. We looked to investigate this interaction from both applied and basic science perspectives. For the applied study, we examined two culvert replacements designed to mimic natural stream channels, to see if they were better at maintaining ecosystem processes within as well as upstream and downstream of culverts compared to non-replaced culverts. We measured three ecosystem processes (nutrient uptake, hydrologic characteristics, and coarse particulate organic matter retention) and found that stream simulation culvert restoration improved organic matter retention within culverts, and that there were no differences in processes measured upstream and downstream of both restoration designs. Our results suggest that measurements of ecosystem processes are more likely to show a response to restoration if they match the scale of the restoration activity. For the basic science study, we quantified the longitudinal spatial heterogeneity of physical and biofilm characteristics at microhabitat to segment scales on streams with different streambed variability. We found that all physical characteristics and biofilm characteristics were spatially independent at the macro-habitat scale and greater. Together, these studies demonstrate the importance of scale in ecological interactions and the value of incorporating considerations of scale into restoration activities.

Investigating within-canopy variation of functional traits and cellular structure of sugar maple (Acer saccharum) leaves

Patterns of increasing leaf mass per area (LMA), area-based leaf nitrogen (Narea), and carbon isotope composition (δ13C) with increasing height in the canopy have been attributed to light gradients or hydraulic limitation in tall trees. Theoretical optimal distributions of LMA and Narea that scale with light maximize canopy photosynthesis; however, sub-optimal distributions are often observed due to hydraulic constraints on leaf development. Using observational, experimental, and modeling approaches, we investigated the response of leaf functional traits (LMA, density, thickness, and leaf nitrogen), leaf carbon isotope composition (δ13C), and cellular structure to light availability, height, and leaf water potential (Ψl) in an Acer saccharum forest to tease apart the influence of light and hydraulic limitations. LMA, leaf and palisade layer thickness, and leaf density were greater at greater light availability but similar heights, highlighting the strong control of light on leaf morphology and cellular structure. Experimental shading decreased both LMA and area-based leaf nitrogen (Narea) and revealed that LMA and Narea were more strongly correlated with height earlier in the growing season and with light later in the growing season. The supply of CO2 to leaves at higher heights appeared to be constrained by stomatal sensitivity to vapor pressure deficit (VPD) or midday leaf water potential, as indicated by increasing δ13C and VPD and decreasing midday Ψl with height. Model simulations showed that daily canopy photosynthesis was biased during the early growing season when seasonality was not accounted for, and was biased throughout the growing season when vertical gradients in LMA and Narea were not accounted for. Overall, our results suggest that leaves acclimate to light soon after leaf expansion, through an accumulation of leaf carbon, thickening of palisade layers and increased LMA, and reduction in stomatal sensitivity to Ψl or VPD. This period of light acclimation in leaves appears to optimize leaf function over time, despite height-related constraints early in the growing season. Our results imply that vertical gradients in leaf functional traits and leaf acclimation to light should be incorporated in canopy function models in order to refine estimates of canopy photosynthesis.

BEYOND ROOTS ALONE: NOVEL METHODOLOGIES FOR ANALYZING COMPLEX SOIL AND MINIRHIZOTRON IMAGERY USING IMAGE PROCESSING AND GIS TOOLS

Quantifying belowground dynamics is critical to our understanding of plant and ecosystem function and belowground carbon cycling, yet currently available tools for complex belowground image analyses are insufficient. We introduce novel techniques combining digital image processing tools and geographic information systems (GIS) analysis to permit semi-automated analysis of complex root and soil dynamics. We illustrate methodologies with imagery from microcosms, minirhizotrons, and a rhizotron, in upland and peatland soils. We provide guidelines for correct image capture, a method that automatically stitches together numerous minirhizotron images into one seamless image, and image analysis using image segmentation and classification in SPRING or change analysis in ArcMap. These methods facilitate spatial and temporal root and soil interaction studies, providing a framework to expand a more comprehensive understanding of belowground dynamics.

THE INFLUENCE OF DIET COMPOSITION, PLANT DEFENSIVE CHEMICALS, AND WINTER SEVERITY ON THE NUTRITIONAL CONDITION OF A FREERANGING, GENERALIST HERBIVORE

Herbivory requires animals to manage intake of toxic phytochemicals. Detoxification and excretion of these chemicals prevents toxicity, but is energetically expensive. I investigated the relationship between investment in detoxification and nutritional condition for moose on Isle Royale National Park (Alces alces) during winter, using urinary indices from urine samples collected in snow. The ratio of urinary urea nitrogen:creatinine is an indicator of nutritional condition, and the ratio of glucuronic acid:creatinine is an indicator of investment in detoxification. Nutritional condition declined with greater investment in detoxification. An alternative means of managing defensive chemical intake is to diversify the diet. Microhistological analysis of fecal pellets determined diet composition. Diet diversity was weakly associated with improved nutritional condition. However, the strongest predictors of nutritional condition were winter severity and proportion of balsam fir in the diet (a dominant food for moose in this ecosystem).

Spatial and temporal patterns in ungulate-ecosystem interactions

Ungulates are important components of a variety of ecosystems worldwide. This dissertation integrates aspects of ungulate and forest ecology to increase our understanding of how they work together in ways that are of interest to natural resource managers, educators, and those who are simply curious about nature. Although animal ecology and ecosystem ecology are often studied separately, one of the general goals of this dissertation is to examine how they interact across spatial and temporal scales. Forest ecosystems are heterogeneous across a range of scales. Spatial and temporal habitat use patterns of forest ungulates tend to be congregated in patches where food and/or cover are readily available. Ungulates interact with ecosystem processes by selectively foraging on plants and excreting waste products in concentrated patches. Positive feedbacks may develop where these activities increase the value of habitat through soil fertilization or the alteration of plant chemistry and architecture. Heterogeneity in ecosystem processes and plant community structure, observed at both stand and local scales, may be the integrated outcome of feedbacks between ungulate behavior and abiotic resource gradients. The first chapter of this dissertation briefly discusses pertinent background information on ungulate ecology, with a focus on white-tailed deer (Odocoileus virginianus) in the Upper Great Lakes region and moose (Alces acles) in Isle Royale National Park, Michigan, USA. The second chapter demonstrates why ecological context is important for studying ungulate ecology in forest ecosystems. Excluding deer from eastern hemlock (Tsuga canadensis) stands, which deer use primarily as winter cover, resulted in less spatial complexity in soil reactive nitrogen and greater complexity in diffuse light compared to unfenced stands. The spatial patterning of herbaceous-layer cover was more similar to nitrogen where deer were present, and was a combination of nitrogen and light within deer exclosures. This relationship depends on the seasonal timing of deer habitat use because deer fertilize the soil during winter, but leave during the growing season. The third chapter draws upon an eight-year, 39-stand data set of deer fecal pellet counts in hemlock stands to estimate the amount of nitrogen that deer are depositing in hemlock stands each winter. In stands of high winter deer use, deer-excreted nitrogen inputs consistently exceeded those of atmospheric deposition at the stand scale. At the neighborhood scale, deer-excreted nitrogen was often in excess of atmospheric deposition due to the patchy distribution of deer habitat use. Spatial patterns in habitat use were consistent over the eight-year study at both stand and neighborhood scales. The fourth chapter explores how foraging selectivity by moose interacts with an abiotic resource gradient to influence forest structure and composition. Soil depth on Isle Royale varies from east to west according to glacial history. Fir saplings growing in deeper soils on the west side are generally more palatable forage for moose (lower foliar C:N) than those growing in shallower soils on the east side. Therefore, saplings growing in better conditions are less likely to reach the canopy due to moose browsing, and fir is a smaller overstory component on the west side. Lastly, chapter five focuses on issues surrounding eastern hemlock regeneration failure, which is a habitat type that is important to many wildlife species. Increasing hemlock on the landscape is complicated by several factors including disturbance regime and climate change, in addition to the influence of deer.

Spatial Ecology of the Great Basin Rattlesnake on the Snake River Plain

Dr. Charles Peterson, Herpetologist and Professor of Biological Sciences at Idaho State University. Curator of Ichthyology and Herpetology at the Idaho Museum of Natural History.

Revegetation and Bird Trends, Silver Bow Creek Superfund Remediation and Restoration

Silver Bow Creek runs approximately 25 miles from Butte to Warm Springs, where it joins Warm Springs Creek to form the Clark Fork River. This historic creek was terribly contaminated with mine wastes around the turn of the 20th century, leaving many "slickens" that persisted into the 21st century, when it became a Superfund remediation project. More than 5.5 million cubic yards of stream-deposited mine waste have been removed and 1,650 acres revegetated. Chief contaminants are copper, zinc, and arsenic, but acidic soils are often equally or more limiting to plants. The stream was relocated, and mine wastes were replaced with biologically inert cover soil. Richard A. Prodgers is currently a plant ecologist with Bighorn Environmental Sciences in Dillon, Montana.

Environmental and Endogenous Factors Influencing Emigration in Juvenile Anadromous Alewives

We analyzed juvenile anadromous alewife migration at Bride Lake, a coastal lake in Connecticut, during summer 2006 and found that migration on 24-hour and seasonal timescales was influenced by conditions of the environment and characteristics of the individual. To identify environmental cues of juvenile migration, we continuously video recorded fish at the lake outflow and employed information-theoretic model selection to identify the best predictors of daily migration rate. More than 80% of the approximately 320,000 juveniles that migrated from mid-June to mid-August departed in three pulses lasting one or two days. Pulses of migration were associated with precipitation events, transient decreases in water temperature and transient increases in stream discharge. Diel timing of migration shifted over the summer. Early in the season most migration occurred around dawn; late in the season migration occurred at night. To identify individual characteristics associated with migratory behavior, we compared migrating juveniles that we collected as they were exiting Bride Lake to non-migrating juveniles that we collected from the center of the lake. Migrants were a non-random subset of the population; they were on average 1 – 12 mm larger, 2 – 14 d older, had grown more rapidly (11% greater length-at-age), and were in better condition (14% greater mass-at-length) than non-migrant fish. We infer that the amount of accumulated energy has a positive effect on the net benefit of migration at any time in the migratory season.

Anadromous Rainbow Smelt and Tomcod in Connecticut: Assessment of populations, conservation status, and need for restoration plan

(beginning of rainbow smelt executive summary) Evidence indicates that anadromous rainbow smelt (Osmerus mordax) populations in Connecticut and elsewhere in the northeast United States have severely declined. Several sampling programs have documented declines in Connecticut’s smelt populations over the last three decades (Marcy 1976a, Marcy 1976b, Millstone Environmental Laboratory 2005). Similar declines have also been documented in the Hudson River (ASA Analysis & Communication 2005) and in Massachusetts (personal communication, Brad Chase, MA Division of Marine Fisheries 2004). Recreational and commercial fisheries in the region for this species have virtually ceased (Blake and Smith 1984). The Connecticut Fish Advisory Committee of the Endangered Species Program has recommended that rainbow smelt be listed as threatened in Connecticut, and the National Marine Fisheries Service (2004) has recently listed rainbow smelt as a Federal Species of Concern. The purpose of this project is to develop an environmental history of rainbow smelt in Connecticut and surrounding regions, and document the current status of populations in Connecticut waters. An environmental history that assesses trends in abundance, environmental threats and historical efforts to ameliorate the threats will contribute to regional efforts to conserve these fish. Comprehensive review of the regional literature and trends associated with rainbow smelt has not been undertaken since Kendall (1926). Assessment of current abundance, distribution, areas of critical habitat, and whether the species is presently reproducing in state waters is critical for clarifying conservation status, designing a monitoring program and developing a recovery or enhancement plan, if this appears to be necessary. (beginning of tomcod executive summary) Atlantic tomcod (Microgadus tomcod) are believed to have declined significantly in Connecticut and other estuaries of the Northeast and Middle Atlantic states. Several monitoring programs indicate that the species is scarce and/or declining in the region’s estuaries (Gottschall and Pacileo 2004, Molnar 2004, Millstone Environmental Laboratory 2005, ASA Analysis and Communication 2005). Once-active recreational (NMFS MRFSS 2005, http://www.st.nmfs.gov) and commercial fisheries for this species in Connecticut are now dormant. For the past 10 years, the Connecticut Fish Advisory Committee of the Endangered Species Program has recommended that studies be undertaken to quantify the status of tomcod populations and to determine if conservation actions should be initiated. The purpose of this project is to develop an environmental history of Atlantic tomcod in Connecticut and surrounding regions, and document the current status of populations in Connecticut waters. An environmental history that assesses trends in abundance, environmental threats and historical efforts to ameliorate the threats will contribute to regional efforts to conserve these fish. Assessment of current abundance, distribution, areas of critical habitat, and whether the species is presently reproducing in state waters is critical for determining conservation status, designing a monitoring program and developing a recovery or enhancement plan, if this appears to be necessary.