Lisa Boudeman Dr.

Bruns Report 3 7 December 2011

Upland Boreal Forest Soils Overlying Oil Sands in Alberta Canada As easily accessible oil deposits around the world diminish, commercial mining of unconventional sources such as bitumen deposits in the boreal Athabasca oil sands region in Alberta, Canada have become an increasingly profitable venture. Although unconventional energy extraction methods increase the worlds oil supplies, they also come with a greater environmental cost than traditional extraction methods (Rooney and Bayley, 2010). Beginning in 1967, large scale commercial oil sands mining began to disrupt the native ecosystem in the boreal forest of Alberta, Canada. In recent years, however, the environmental impact of oil extraction from tar sands, particularly in Canada, has become a topic of interest in the science community. Extracting the bitumen from these sands often requires stripping the overlying forest, peat, and soil of entire landscapes, and digging up to one hundred meters below the surface, leaving barren landscapes dominated by overburden dumps, toxic tailings impoundments, and open mine pits (Trites and Bayley, 2009). Currently, more than 60,200 ha of land area has been disturbed an additional 480,000 ha are expected to be mined (Rooney and Bayley, 2011). Extracting the estimated 1.6 trillion barrels of bitumen found in the oil sands in Alberta will require stripping 2000 km2 of boreal forest (Dimitriu et. al, 2010). Due to the destructive nature of this process, the land cannot be bioremediated, but instead, must be physically reclaimed. Prior to oil sands mining, wetlands and, to a lesser extent, upland forest habitats dominated the Athabasca boreal landscape. Due to the difficultly in wetland reclamation, nearly half of the area once dominated by wetlands will be reclaimed to mimic native upland soils such as the trembling aspen, balsam poplar, and white spruce stand located in Breton, Alberta (Trites and Bayley, 2009). To understand the great complexity of recreating a native upland soil after oil mining, one first must examine the geology and extensive soil ecological interactions of a true native soil. Although an assortment of tree stands exist in the upland boreal forest region that overlies the oil sand deposits, a trembling aspen, balsam poplar, and white spruce stand such as that in Breton, Alberta represents a typical boreal forest upland (Dimitriu, et al., 2010). At the foundation of this native upland soil lies a glacial till parent material first noted at a depth of 81 cm. This particular site is well-drained with the water table falling below 4.25 meters (Howitt and Pawluk, 1985). The mixed deciduous-conifer overlying stand structure allows for a diverse set of habitats for soil biota in this region. In the deciduous dominated areas, the cool temperature regime enhances an accumulation of aspen and poplar leaves. The dynamic nature of leaf material accumulation prohibits the development of slow growing organisms such as bryophytes and lichens under deciduous trees (Crites et al., 1998). However, the thick cover of leaf material provides a substantial amount of cool, moist, protected habitat and low-lignin, calcium and nutrient rich food for many organisms such as microarthropods, mites, heterotrophic microbes, and fungi to thrive (Laganiere et al., 2009). Underlying the coniferous vegetation, in contrast, ground exposure provides a consistent habitat for organisms that are sensitive to environmental changes such as lichens and mosses. Downed woody material (DWM) provides a third significant habitat source for soil organisms, providing a stable, nutrient rich environment for the proliferation of lichens, fungal mycelia, chitin-degrading bacteria (actinomycetes), and microfungi, as well as a protected space for microarthropods (Crites et al., 1998). Finally, the fine to medium sized roots of the macrophytes, (ie. the rhizosphere), provides an excellent source of nutrients and habitat for organisms such as mycorrhizal fungi. As a result of the
1

Lisa Boudeman Dr. Bruns Report 3 7 December 2011

aforementioned soil forming factors acting on the soil here (parent material, climate, landscape position, biological factors, and time), this upland soil has been designated as a gray luvisol (known as a boralf in the U.S. taxonomy system) (Agriculture and Agri-Food Canada, 2011). A substantial dynamic organic layer dominated by aspen and poplar leaves underlies the predominantly deciduous portions of this stand. The most superficial layer, the Oi horizon, is approximately 3 cm thick, consisting of non-decomposed and semi-decomposed spruce needles and aspen leaves (Howitt and Pawluk, 1985). In this zone, microarthropods play a significant role in the soil ecosystem by grazing on coarse leaf material, a process that increases the litter surface area and releases and distributes nutrients for microbes and fungi (Visser et al., 1987). The next organic layer, the Oe horizon, is approximately 4 cm thick and consists of some decomposed leaves, needles, and an abundant amount of fine, very fine, and medium sized roots (Howitt and Pawluk, 1985). Here, microarthropods thrive off of leaf and root organic matter, playing a key role in regulating rates of decomposition and nutrient mineralization, the distribution of organic nutrients, and the formation of stable soil structure and humus. One group of soil microarthropods of particular interest, the Onychiuridae (a collembolan family) clearly plays a vital role in nutrient cycling in these soils. In general, collembola, a group of arthropods that number up to 100,000 individuals per m2 of soil, transport nutrients, microbes, mycoparasites (fungal parasites), saprophytes (which contain decay enhancing chitinases), and fungal spores (which can colonize new areas) to various soil and organic layers via fecal and skin deposition as a result of vertical channeling (Visser et al., 1987). Collembola are distinguished by their ventral tube (collophore), an organ that senses the moisture status of the soil substrate and can absorb liquids, ions, and gases, both processes which are essential in collembolan osmoregulation and in preventing desiccation (Coleman et al., 2004). The ventral tube also can function as a sticky appendage to aid in attaching to slippery surfaces (Hopkin, 2002). Collembola have three body regions, one pair of antennae, and are classified as hexapods (mandibulate arthropods with six legs). The Onychiuridae family is typically podomorphic and euedaphic or hemiedaphic, that is, they live in the lower litter layers and upper mineral layers of the soil. To adapt to this habitat, the Onychiuridae lack a furcula (the jumping organ typical of edaphic collembola), eyes, and pigmentation, and have developed psudocelli, a cuticular organ that can extrude defensive oils when disturbed (Coleman et al., 2004). In the soil, collembola aggregate not only as a result of habitat conditions and food availability, but also as a behavioral response to other collembola pheromonessignals which can attract the opposite sex for reproduction. Although some collembola demonstrate parthenogenesis, most species reproduce sexually via sperm transfer. In this process, the male deposits his sperm in the form of spermatophores, packets of sperm, either actively onto the female or passively onto pedicels for the female to find (Coleman et al., 2004). Once the eggs are fertilized, they are laid in groups where they will hatch and go through a series of two to 50 instars, a developmental process which can take 40 to 400 days (Lavelle and Spain, 2001). Unlike insects, collembola continue to molt throughout their lifetime. Collembola demonstrate opportunistic growth patternsunder suitable conditions, collembola can grow and reproduce explosively, resulting in collembola densities ranging from 10,000 to 100,000 collembola per m2 (Hopkin, 2002). As noted above, the Onychiuridae collembola grazing and migration patterns significantly contribute to nutrient cycling and microbial dispersal in the boreal forest regions. The Onychiurus subtenuis grazes primarily on pigmented fungi hyphae of fungi in the genus Cladosporium (*Hassall, 1986). Hence, Onychiurus subtenuis are not randomly dispersed
2

Lisa Boudeman Dr. Bruns Report 3 7 December 2011

throughout the bulk soil, but instead, aggregate in colonies 8 to 30 centimeters in diameter surrounding the rhizosphere (the nutrient-rich soil surrounding plant roots) of the plants in association with the Cladosporium (Lavelle and Spain, 2001). In the grazing process, these critters grind large organic particles into smaller pieces, which increases organic matter decomposition and mobilization of nutrients for other organisms by increasing the surface area subject to microbial attack. By increasing decomposition, these critters enhance the formation of the humus horizon, which provides a stabilized carbon source for producers and microbes, and also demonstrates a prime example of collembola as one of the many organisms contributing to the biotic soil forming factors. Not only does Onychiuridae grazing enhance nutrient cycling by microbes, but grazing on fungal mycelia can also increase oxygen uptake by fungal mycelia, stimulate fungal growth, and increase overall fungal activity (Hassall, 1986). Because collembola are moisture dependent, their grazing patterns depend on seasonal and precipitation changes, often resulting in extensive vertical migration. For example, to prevent desiccation in the summertime, Onychiurus subtenuis aggregate in the lower, moist litter (humus) layers surrounding areas of highest fungal mycelia density. After rainfall events, these critters undergo significant vertical migration (potentially 1000 or more per m2) to the superficial litter layers to graze on the abundance of fungal mycelia and microorganisms in this level (*Hassell, 1986). When the Onychiurus subtenuis migrate, they carry spores and fungi on their body surface and in their gut, dispersing thousands of spores and microbes from 100 or more taxa of fungi per m2 throughout the profile as they journey through the soil layers. As a result, Onychiuridae enhance biodiversity by diffusing otherwise immobile microbes and fungal spores to inoculate new nutrient-rich areas. If no other functional species exists in these areas, these inoculants can potentially increase the rate of leaf litter decomposition (Hassall, 1986). Not only does vertical channeling enhance biodiversity and dispersal of fungi and microbes, but this vertical channeling also opens up macropores that allow plant roots to access nutrient and moisture rich spaces that may have previously been inaccessible which may enhance plant growth as well. Additionally, vertical channeling increases mass water flow, an outcome which can enhance dispersal of organisms with little channeling ability that rely on previously created macropores and/or water flow for movement, reproduction, and for access to food sources. After taking an in-depth look into the collembolas extensive interaction web with other organisms and applying this idea to the organisms described throughout this paper, one can see that restoring a soil after complete obliteration from the oil sands mining process may have strong environmental repercussions if biological webs are not considered in the physical reclamation process. In addtiona to collembola, other microarthropods such as orbatid, prostigmatid, and mesostigmatid mites are also key detritivores and fungivores in these soils. The most abundant mite, the orbatid is very sensitive to deforestation, and therefore acts as an indicator of habitat destruction (Lindo and Visser, 2004). In addition to microarthropods, the moist, nutrient rich, more stabilized nature of the Oe layer, promotes the growth an abundance of fungal hyphae from the basidiomycota Piloderma sp. and Russula sp, and saprotrophs such as Hydnellum sp. (Demitriu et al., 2010). The Oa horizon, the oldest, most decomposed organic horizon, consists of decomposed fibrous to matted organic matter; abundant, fine and very fine random and plentiful, medium random roots (Howitt and Pawluk, 1985). Here, the vast quantity of root exudates and highly degraded organic material would promote the growth of organisms such as actinomycetes and aerobic bacteria. In contrast to areas with thick leaf litter accumulation, areas directly beneath coniferous trees or on downed woody material (DWM) promote an enrichment of slow growing, slow
3

Lisa Boudeman Dr. Bruns Report 3 7 December 2011

moving, soil organisms that are sensitive to habitat change. In particular, DWM serves an essential role in nutrient and organic material dynamics in the boreal ecosystem. For example, the accumulation of woody debris promotes the proliferation of chitin-degrading bacteria (actinomycetes) which can break down diverse polymers such as fats and waxes that would otherwise be inaccessible to soil microbes. As a result, DWM supports nutrient cycling in the boreal forest region. DWM also enhances the growth of fungal mycelia of ectomycorrhizal fungi such as Piloderma sp., resulting in enhanced nutrient and water absorption for other macrophytes. Also, DWM promotes diversity, providing a stable superficial substrate for bryophytes, mosses, liverworts, and lichens that would otherwise not be able to inhabit these areas. Due to the sensitive nature of both liverworts and lichens, the presence or absence of these species acts as an important indicator of ecosystem disruption and pollution (Crites et al., 1998). The first mineral layer underlying the mixed deciduous-coniferous stand, identified as Aeh (0-3 cm), is dark grayish brown in color, an indication of illuviation of organic material from the overlying organic layers and an eluviation of clay to subsurface horizons. This horizon is a sandy loam with a fine granular structure and is very friable (the peds break easily under pressure). The illuviation of organic matter from the overlying layers and the porous structure of this layer create a well-aerated, nutrient-rich habitat for the proliferation of fine to medium sized roots (Howitt and Pawluk, 1985). Here, tree roots provide soil structure and excellent habitat and energy source for microarthropods, microbes, and, of great significance, ectomycorrhizal fungi. The symbiotic relationship between ectomycorrhizal fungi and tree roots plays a noteworthy role in maintaining the health of a tree stand. Here, ectomycorrhizal fungi, such as Piloderma sp., colonize fine roots of trees, increasing nutrient absorption, pathogen resistance, and tolerance to drought and heavy metal toxins by providing hyphae extensions into the soil areas that root hairs would typically be unable to access. In turn, the trees provide energy (sugar) for these fungi, thus promoting fungal growth that will aid in tree health (DeBellis et al., 2006). Underlying the Aeh horizon (from 3-8 cm) is a grayish brown Ae horizon with a sandy loam texture; the lighter coloration is indicative of organic matter eluviation to lower horizons. Despite loss of organic material, this horizon still contains plentiful fine and very fine random and horizontal roots (Howitt and Pawluk, 1985). The relatively coarse soil textural class and fine roots provides adequate tunneling room for collembolan and other macroinvertebrates as well as nutrients for microbes and fungi. Of recent interest, the exotic earthworm species Lumbricus rubillus moves through these layers, grazing on the fungi and bacteria the rhizosphere, breaking down large litter pieces, inoculating them with microbial decomposers, and excreting nutrient rich feces into the mineral soil; through this tunneling action, they both facilitate water and gas flux and enhancing nutrient cycling (Laganiere, 2009). Although microarthropods have been shown to enhance community development through their tunneling actions, the Lumbricus rubillus can rapidly remove the forest floor when it invades, which drastically influences the understory plant communities, and, as a result, diminishes the nutrients and shelter availability for other organisms here (National Resources Research Institute, 2011). The Ae2 horizon (8-13 cm) is grayish brown with a loamy sand texture; the change in texture class most likely indicates further elluviation of clays out of the surface horizons. Here, there are few, fine to very fine oblique and random roots that provide nutrients and habitat for soil microbes and mycorrhizal fungi. Although soil roots are still prevalent and the coarse soil texture still promotes macropore formation (facilitating proper aeration and biota movement), a decrease in the amount of decaying organic matter and number of roots to provide nutrients for
4

Lisa Boudeman Dr. Bruns Report 3 7 December 2011

living organisms will result in a significant decrease in soil biota abundance and diversity in this layer, and will continue to decrease with increasing depth. The next horizon, the AB horizon (13-17 cm), is yellowish brown with a loam to sandy clay loam texture. This transitional horizon has properties of both an A and a B horizon, indicated by the combination of a weak, coarse platy to moderate, firm, medium blocky structure. Here, there are few, medium and fine random roots. Underlying the AB horizon is an argillic, Bt horizon (17-23 cm), a horizon designation descriptive of an influx of clay. This horizon is a yellowish brown clay loam that has a strong, fine subangular blocky structure and friable to firm moist consistence. Here, as a result of increased clay content (which equates to fewer macropores) and increased depth beneath the surface, there are only a few, fine random roots and few (if any) macroinvertebrates. The next argillic horizon (23-47 cm), Bt2, is a brown clay loam with strong, fine to medium subangular blocky, and weak, medium prismatic structure. Here, few, fine and very fine random roots persist. The final Bt layer, Bt3 (47-67 cm), has the largest illuviation of clay from overlying layers. Bt3 is a dark yellowish brown-olive brown clay loam, with moderate, medium to coarse blocky and weak, medium prismatic structure. This horizon has a firm moist consistence, and continuous, moderately thick clay films evident along cleavage faces; high clay content and increasing depth hinders biota growth as evidenced by very few medium oblique roots. Beginning at 67 cm, pedogenic processes are still in the early stages. Here, the dark yellowish brown-olive brown BC horizon (67-81 cm) is a sandy clay loam to clay loam with weak, coarse blocky structure and firm moist consistency with fine random roots. Starting at BC2 (81-111 cm), the glacial till parent material becomes apparent. The BC2 horizon is a dark yellowish brown-olive brown sandy clay loam with fragmental glacial till and very few, medium to fine horizontal and vertical roots. The BC3 horizon is marked by an increase in glacial till; this horizon is a light olive brown sandy clay to sandy clay loam till (Howitt and Pawluk, 1985). Since 1967, oil sands mining have resulted in a significant amount of soil destruction in the Athabasca oil sands region similar to the preceding soil ecological description (Trites and Bayley, 2009). Due to an increased demand for the worlds limited oil supply, extracting oil from unconventional supplies such as Athabasca oil sands has gained great economic appeal. As noted at the beginning of this paper, surface and subsurface soils as well as all overlying vegetation must be completely removed in order to extract oil from these sands. Aside from the initial destruction of these diverse habitats, the reclamation of these lands may pose just as much impact on these communities. Not only are half of the native wetlands converted to uplands, but also these reclaimed areas have reduced horizon stratification, excessive amounts of salts and contaminants, and altered nutrient availability relative to native upland areas. Where pre-disturbed upland and wetlands have significant stratification and profile development, reclaimed soils are composed of two relatively homogenous layers: a surface layer composed of a mineral material/peat mixture and a deep layer composed of tailings sand and overburden material or geological parent material. In reclaimed lands where tailings sand is used, the soil has increased salinity, residual bitumen, and hydrocarbon toxicants, each of which pose serious hurdles to the reestablishment of the native soil communities (Rooney and Bayley, 2011). In addition to limiting toxicants, revegetation practices have hindered native succession. As described above, the native fungi, microbes, and microarthropods of this region interact closely with the tree stands in the boreal region and depend on them for nutrients and shelter which are provided by their roots, leaves, and DWM. Not only do these organisms depend on the tree stands, but these soil organisms also depend on each other through a tightly integrated soil web. For example, these complex relations can be seen the grazing and migration patterns of simply
5

Lisa Boudeman Dr. Bruns Report 3 7 December 2011

one collembola family, the Onychiuridae; the many roles of this family include dispersing fungal spores and microbes, shredding organic matter which increases decomposition rates, humus formation, and nutrient mineralization for other organisms, enhancing fungal hyphae growth through grazing, and tunneling which enhances root growth and increases mobility for organisms with limited mobility. Unfortunately, these intricate soil ecological interactions fly under the radar in the reconstruction of the Athabasca region following oil sands mining. In typical reclaimed areas, revegetation often includes planting barley which favors the growth of arbuscular mycorrhizae such as Preussia sp. and Giberella sp. and restricts ectomycorrhizal growth typical of the boreal forest region. As demonstrated by Dimitriu et al., however, even after forest vegetation recuperates, the vast alteration in abiotic soil factors continues to hinder, if not completely prevent, the succession of native soil biota (2010). In general, reclaimed lands continue to have biota vastly differing from the native populations and support lower enzyme activities, more grass and forbs, higher bacteria:fungi ratios, higher pH and more endophytic fungi (Dimitriu et al., 2010). Before oil sands mining operations destroys the remaining native boreal habitats in the Athabasca region in its entirety, further study on ecosystem dynamics will be imperative to better understand the needs for this biological network to flourish in reclaimed lands. Ideally, once additional knowledge of this soil system is uncovered, the original native ecological systems will be proficiently reestablished from the barren landscapes left behind by oil sands operations.

Lisa Boudeman Dr. Bruns Report 3 7 December 2011

Works Cited Agriculture and Agri-Food Canada. Chapter 16: Correlation of Canadian Soil Taxonomy with Other Systems. The Canadian System of Soil Classification (Third Edition). Web. 11 June 2010. 21 October 2011. <http://sis.agr.gc.ca/cansis/taxa/cssc3/chpt16.html>. Coleman et al. Collembola. Fundamentals of Soil Ecology. 2nd ed. Burlington: Elsevier Academic Press, 2004. 101-109. Crites, Susan et al. Diversify and abundance of bryophytes, lichens, and fungi in relation to woody substrate and successional stage in aspen mixedwood boreal forests. Canadian Journal of Botany. 76 (1998): 641-651. DeBellis, T. et al. Relationships between Stand Composition and Ectomycorrhizal Community Structure in Boreal Mixed-Wood Forests. Microbial Ecology. 52 (2006): 114-126. Dimitriu, Pedro A. et. al. Impact of reclamation of surface-mined boreal forest soils on microbial community composition and function. Soil Biology and Biochemistry 42 (2010): 2289-2297. Howitt, R. W. and S. Pawluk. The Genesis of a Gray Luvisol within the Boreal Forest Region. I. Static Pedology. Canadian Journal of Soil Science. 65:1 (1985): 1-8. Hassall, M. Effects of the collembolan Onychiurus subtenuis on decomposition of Populus tremuloids leaf litter. Pedobiologa. 29 (1986): 219-225. *Hassall. M. Vertical Migration of Onychiurus-subtenuis Collembola in Relation to Rainfall and Microbial Activity. Pedobiologa. 29 (1986): 175-192. Hopkin, Steve P. Collembola. Encyclopedia of Soil Science. United Kingdom: Markel Dekker Inc, 2002. 207-210. Lavelle, Patrick. Spain, Alister V. Collembola. Soil Ecology. Netherlands: Kluwer Acadimic Publishers,2001. 263-270. Laganiere, Jerome et al. Linking the abundance of aspen with soil faunal communities and rates of belowground processes within single stands of mixed aspen-black spruce. Applied Soil Ecology. 41 (2009): 19-28. Lindo, Zoe and Suzanne Visser. Forest floor microarthropod abundance and oribatid mite (Acari: Oribatida) composition following partial and clear-cut harvesting in the mixedwood boreal forest. Canadian Journal of Forest Resources. 34 (2004): 998-1006. Natural Resources Research Institute. Worm Identification - Earthworm Ecological Groups. 2011. 27 October 2011. <http://www.nrri.umn.edu/worms/identification/ecology_groups.html>.

Lisa Boudeman Dr. Bruns Report 3 7 December 2011

Rooney, Rebecca C. and Suzanne E. Bayley. Setting reclamation targets and evaluating progress: Submersed aquatic vegetation in natural and post-oil sands mining wetlands in Alberta, Canada. Ecological Engineering 37 (2011): 569-579. Trites, Marsha and Suzanne E. Bayley. Vegetation communities in continental boreal wetlands along a salinity gradient: Implications for oil sands mining reclamation. Aquatic Botany 91 (2009): 27-39. Visser, Suzanne et. al. Fungi associated with Onychiurus subtenuis (Collembola) in an aspen woodland. Canadian Journal of Botany. 65 (1987): 635-642.