Ecological dynamics
The information in this Ecological Site Description, including the state-and-transition model (STM), was developed based on historical data, current field data, professional experience, and a review of the scientific literature. As a result, all possible scenarios or plant species may not be included. Key indicator plant species, disturbances, and ecological processes are described to inform land management decisions.
The MLRA lies within the tallgrass prairie ecosystem of the Midwest. The heterogeneous topography of the area results in variable microclimates and fuel matrices that in support prairies, savannas, and forests. Loess Upland Forests form an aspect of this vegetative continuum. This ecological site occurs on uplands on well-drained soils. Species characteristic of this ecological site include a closed canopy of oaks with shade-tolerant herbaceous vegetation.
Fire is a critical factor that maintains Loess Upland Forests. Fire typically consisted of low-severity surface fires every 25 to 50 years (LANDFIRE 2009). Ignition sources included summertime lightning strikes from convective storms and bimodal, human ignitions during the spring and fall seasons. Native Americans regularly set fires to improve sight lines for hunting, drive large game, improve grazing and browsing habitat, agricultural clearing, and enhance vital ethnobotanical plants (Barrett 1980; LANDFIRE 2009).
Drought and storm damage have also played a role in shaping this ecological site. The periodic episodes of reduced soil moisture in conjunction with the well-drained soils have favored the proliferation of plant species tolerant of such conditions. Drought can also slow the growth of plants and result in dieback of certain species. Damage to trees from wind and ice storms can vary from minor, patchy effects of individual trees to stand effects that temporarily affect community structure and species richness and diversity (Irland 2000; Peterson 2000). When coupled with fire, periods of drought and catastrophic storm damage can greatly delay the establishment and maturation of woody vegetation (Pyne et al. 1996).
Today, Loess Upland Forests have been reduced from their pre-settlement extent. Low to moderate slopes have been converted to cropland, while steeper slopes have been converted to forage land. Remnants that do exist have experienced long-term fire suppression and overbrowsing resulting in significant changes to the forest structure. A return to the historic plant community may not be possible following extensive land modification, but long-term conservation agriculture or forest reconstruction efforts can help to restore some biotic diversity and ecological function. The state-and-transition model that follows provides a detailed description of each state, community phase, pathway, and transition. This model is based on available experimental research, field observations, literature reviews, professional consensus, and interpretations.
State 1
Reference State
The reference plant community is categorized as an oak forest, dominated by deciduous trees and shade-tolerant herbaceous vegetation. The one community phase within the reference state is dependent on recurring fire intervals. The severity and intensity of fire alters species composition, cover, and extent, while regular fire intervals keep the canopy from succeeding to mesophytic, fire-intolerant species. Drought and catastrophic storm damage have more localized impacts in the reference phases, but do contribute to overall species composition, diversity, cover, and productivity.
Community 1.1
White Oak – Black Oak/Hophornbeam – American Hazelnut/American Hogpeanut – Feathery False Lily of the Valley
White Oak – Black Oak/Hophornbeam – American Hazelnut/American Hogpeanut – Feathery False Lily of the Valley – Sites in this reference community phase are a closed canopy forest. White oak and black oak are the dominant species, but bur oak, northern red oak, and shagbark hickory are common canopy associates. Trees are large (21 to 33-inch DBH), and cover is approximately 80 percent (LANDFIRE 2009). Hophornbeam is regularly found in the subcanopy, and tall shrubs – e.g., American hazelnut – can be present. The herbaceous layer is nearly continuous with shade-tolerant species such as American hogpeanut, feathery false lily of the valley, pointedleaf ticktrefoil (Desmodium glutinosum (Muhl. ex Willd.) Alph. Wood), and spotted geranium (Geranium maculatum L.) (NatureServe 2018). Spring ephemerals, such as mayapple (Podophyllum peltatum L.), bloodroot (Sanguinaria canadensis L.), and Virginia springbeauty (Claytonia virginica L.) can be very abundant in the early spring before the trees have leafed out. Low-severity surface fires every 25 to 50 years will maintain this community phase.
State 2
Fire Suppressed State
Fire suppression can transition the reference plant community from an oak forest to an oak-maple mesophytic forest. As the natural fire regime is removed from the landscape, encroachment and dominance by shade-tolerant, fire-intolerant species ensues. This results in a positive feedback loop of mesophication whereby plant community succession continuously creates cool, damp shaded conditions that perpetuate a closed canopy ecosystem (Nowacki and Abrams 2008). Succession to this forested state can occur in as little as 50 years from the last fire (LANDFIRE 2009). Overbrowsing by an unnaturally abundant deer population can also lead to changes in the composition, diversity, and production of the forest. Continuous browsing has been reported to prevent the regeneration of the historic canopy, which is replaced by mid-level and invasive species (Gubanyi et al. 2008; VerCauteren and Hygnstrom 2011). Similarly, herbaceous diversity and composition is also affected by selective browsing pressure (Gubanyi et al. 2008).
Community 2.1
White Oak – Sugar Maple/Honeysuckle – Common Buckthorn/Virginia Springbeauty – Mayapple
White Oak – Sugar Maple/Honeysuckle – Common Buckthorn/Virginia Springbeauty – Mayapple – This community phase represents the early stages of long-term fire suppression and overbrowsing. Mature white and black oaks are still present, but the more shade tolerant sugar maple (Acer saccharum Marshall) begins to co-dominate. The tree canopy closes to 100 percent cover and basal area increases (LANDFIRE 2009). Non-native shrubs, such as honeysuckle (Lonicera L.) and common buckthorn (Rhamnus cathartica L.), can rapidly colonize. The herbaceous layer continues to support shade-tolerant species, but diversity is reduced as the fully closed canopy results in favorable conditions mostly by spring ephemerals. Grazing pressure alters species composition, allowing plants such as mayapple to increase as it is commonly avoided by deer (Gubanyi et al. 2008; Rawbinski 2008).
Community 2.2
Sugar Maple/Honeysuckle – Common Buckthorn/Mayapple – Litter
Sites falling into this community phase have a well-established, fire-intolerant canopy dominated by sugar maple. Oak seedlings are virtually absent from the understory due to the lack of available light. Without recurring fire, downed woody debris and leaf litter are frequently encountered on the forest floor.
Pathway 2.1A
Community 2.1 to 2.2
Continued fire suppression and increasing deer populations
Pathway 2.2A
Community 2.2 to 2.1
Increased fire intervals, decreased deer populations
State 3
Forage State
The forage state occurs when the reference state is converted to a farming system that emphasizes domestic livestock production known as grassland agriculture. Fire suppression, periodic cultural treatments (e.g., clipping, drainage, soil amendment applications, planting new species and/or cultivars, mechanical harvesting) and grazing by domesticated livestock transition and maintain this state (USDA-NRCS 2003). Early settlers seeded non-native species, such as smooth brome (Bromus inermis Leyss.) and Kentucky bluegrass (Poa pratensis L.), to help extend the grazing season. Over time, as lands were continuously harvested or grazed by herds of cattle, the non-native species were able to spread and expand across the landscape, reducing the native species diversity and ecological function.
Community 3.1
Hayfield
Hayfield – Sites in this community phase consist of forage plants that are planted and mechanically harvested. Mechanical harvesting removes much of the aboveground biomass and nutrients that feed the soil microorganisms (Franzluebbers et al. 2000; USDA-NRCS 2003). As a result, soil biology is reduced leading to decreases in nutrient uptake by plants, soil organic matter, and soil aggregation. Frequent biomass removal can also reduce the site’s carbon sequestration capacity (Skinner 2008).
Community 3.2
Continuous Pastured Grazing System
Continuous Pastured Grazing System – This community phase is characterized by continuous grazing where domestic livestock graze a pasture for the entire season. Depending on stocking density, this can result in lower forage quality and productivity, weed invasions, and uneven pasture use. Continuous grazing can also increase the amount of bare ground and erosion and reduce soil organic matter, cation exchange capacity, water-holding capacity, and nutrient availability and retention (Bharati et al. 2002; Leake et al. 2004; Teague et al. 2011). Smooth brome, Kentucky bluegrass, and white clover (Trifolium repens L.) are common pasture species used in this phase. Their tolerance to continuous grazing has allowed these species to dominate, sometimes completely excluding the native vegetation.
Community 3.3
Rest-Rotation Pastured Grazing System
Rest-Rotation Pastured Grazing System – This community phase is characterized by rotational grazing where the pasture has been subdivided into several smaller paddocks. Through the development of a grazing plan, livestock utilize one or a few paddocks, while the remaining area is rested allowing plants to restore vigor and energy reserves, deepen root systems, develop seeds, as well as allow seedling establishment (Undersander et al. 2002; USDA-NRCS 2003). Rest-rotation pastured grazing systems include deferred rotation, rest rotation, high intensity – low frequency, and short duration methods. Vegetation is generally more diverse and can include orchardgrass (Dactylis glomerata L.), timothy (Phleum pretense L.), red clover (Trifolium pratense L.), and alfalfa (Medicago sativa L.). The addition of native prairie species can further bolster plant diversity and, in turn, soil function. This community phase promotes numerous ecosystem benefits including increasing biodiversity, preventing soil erosion, maintaining and enhancing soil quality, sequestering atmospheric carbon, and improving water yield and quality (USDA-NRCS 2003).
Pathway 3.1A
Community 3.1 to 3.2
Mechanical harvesting is replaced with domestic livestock and continuous grazing
Pathway 3.1B
Community 3.1 to 3.3
Mechanical harvesting is replaced with domestic livestock and rest-rotational grazing
Pathway 3.2A
Community 3.2 to 3.1
Domestic livestock is replaced by mechanical harvesting
Pathway 3.2B
Community 3.2 to 3.3
Implementation of rest-rotational grazing
Pathway 3.3B
Community 3.3 to 3.1
Domestic livestock grazing is replaced by mechanical harvesting
Pathway 3.3A
Community 3.3 to 3.2
Implementation of continuous grazing
State 4
Cropland State
The continuous use of tillage, row-crop planting, and chemicals (i.e., herbicides, fertilizers, etc.) has effectively eliminated the reference community and many of its natural ecological functions in favor of crop production. Corn and soybeans are the dominant crops for the site, and oats (Avena L.) and alfalfa (Medicago sativa L.) may be rotated periodically. These areas are likely to remain in crop production for the foreseeable future.
Community 4.1
Conventional Tillage Field
Conventional Tillage Field – Sites in this community phase typically consist of monoculture row-cropping maintained by conventional tillage practices. They are cropped in either continuous corn or corn-soybean rotations. The frequent use of deep tillage, low crop diversity, and bare soil conditions during the non-growing season negatively impacts soil health. Under these practices, soil aggregation is reduced or destroyed, soil organic matter is reduced, erosion and runoff are increased, and infiltration is decreased, which can ultimately lead to undesirable changes in the hydrology of the watershed (Tomer et al. 2005).
Community 4.2
Conservation Tillage Field
Conservation Tillage Field – This community phase is characterized by rotational crop production that utilizes various conservation tillage methods to promote soil health and reduce erosion. Conservation tillage methods include strip-till, ridge-till, vertical-till, or no-till planting systems. Strip-till keeps seedbed preparation to narrow bands less than one-third the width of the row where crop residue and soil consolidation are left undisturbed in-between seedbed areas. Strip-till planting may be completed in the fall and nutrient application either occurs simultaneously or at the time of planting. Ridge-till uses specialized equipment to create ridges in the seedbed and vegetative residue is left on the surface in between the ridges. Weeds are controlled with herbicides and/or cultivation, seedbed ridges are rebuilt during cultivation, and soils are left undisturbed from harvest to planting. Vertical-till systems employ machinery that lightly tills the soil and cuts up crop residue, mixing some of the residue into the top few inches of the soil while leaving a large portion on the surface. No-till management is the most conservative, disturbing soils only at the time of planting and fertilizer application. Compared to conventional tillage systems, conservation tillage methods can improve soil ecosystem function by reducing soil erosion, increasing organic matter and water availability, improving water quality, and reducing soil compaction.
Community 4.3
Conservation Tillage Field/Alternative Crop Field
Conservation Tillage Field/Alternative Crop Field – This community phase applies conservation tillage methods as described above as well as adds cover crop practices. Cover crops typically include nitrogen-fixing species (e.g., legumes), small grains (e.g., rye, wheat, oats), or forage covers (e.g., turnips, radishes, rapeseed). The addition of cover crops not only adds plant diversity but also promotes soil health by reducing soil erosion, limiting nitrogen leaching, suppressing weeds, increasing soil organic matter, and improving the overall soil ecosystem. In the case of small grain cover crops, surface cover and water infiltration are increased, while forage covers can be used to graze livestock or support local wildlife. Of the three community phases for this state, this phase promotes the greatest soil sustainability and improves ecological functioning within a cropland system.
Pathway 4.1A
Community 4.1 to 4.2
Less tillage, residue management
Pathway 4.1B
Community 4.1 to 4.3
Less tillage, residue management and implementation of cover cropping
Pathway 4.2A
Community 4.2 to 4.1
Intensive tillage, remove residue and reinitialize monoculture row cropping
Pathway 4.2B
Community 4.2 to 4.3
Implementation of cover cropping
Pathway 4.3B
Community 4.3 to 4.1
Intensive tillage, remove residue and reinitialize monoculture row cropping
Pathway 4.3A
Community 4.3 to 4.2
Remove cover cropping
State 5
Reconstructed Oak Forest State
The combination of natural and anthropogenic disturbances occurring today has resulted in numerous forest health issues, and restoration back to the historic reference condition may not be possible. Forests are being stressed by non-native diseases and pests, habitat fragmentation, changes in soil conditions, and overabundant deer populations on top of naturally-occurring disturbances (severe weather and native pests) (IFDC 2018). However, these habitats provide multiple ecosystem services including carbon sequestration; clean air and water; soil conservation; biodiversity support; wildlife habitat; timber, fiber, and fuel products; as well as a variety of cultural activities (e.g., hiking, camping, hunting) (Millennium Ecosystem Assessment 2005; IFDC 2018). Therefore, conservation of forests and woodlands should still be pursued. Forest reconstructions are an important tool for repairing natural ecological functioning and providing habitat protection for numerous species associated with Loess Upland Forests. Therefore, ecological restoration should aim to aid the recovery of degraded, damaged, or destroyed ecosystems. A successful restoration will have the ability to structurally and functionally sustain itself, demonstrate resilience to the ranges of stress and disturbance, and create and maintain positive biotic and abiotic interactions (SER 2002). The reconstructed oak forest state is the result of a long-term commitment involving a multi-step, adaptive management process.
Community 5.1
Early Successional Reconstructed Forest
Early Successional Reconstructed Forest – This community phase represents the early community assembly from forest reconstruction. It is highly dependent on the current condition of the site based on past and current land management actions, invasive species, and proximity to land populated with non-native pests and diseases. Therefore, no two sites will have the same early successional composition. Technical forestry assistance should be sought to develop suitable conservation management plans.
Community 5.2
Late Successional Reconstructed Forest
Late Successional Reconstructed Forest – Appropriately timed management practices (e.g., prescribed fire, hazardous fuels management, forest stand improvement, continuing integrated pest management) applied to the early successional community phase can help increase the stand maturity, pushing the site into a late successional community phase over time. A late successional reconstructed forest will have an uneven-aged canopy and a well-developed shrub layer and understory.
Pathway 5.1A
Community 5.1 to 5.2
Invasive species control and implementation of disturbance regimes
Pathway 5.2A
Community 5.2 to 5.1
Drought or improper timing/use of management actions
Transition T1A
State 1 to 2
Long-term fire suppression and/or land abandonment
Transition T1B
State 1 to 3
Cultural treatments are implemented to increase forage quality and yield
Transition T1C
State 1 to 4
Agricultural conversion via tillage, seeding and non-selective herbicide
Transition T2A
State 2 to 3
Cultural treatments are implemented to increase forage quality and yield
Transition T2B
State 2 to 4
Agricultural conversion via tillage, seeding and non-selective herbicides
Transition R2A
State 2 to 5
Site preparation, tree planting, non-native species control and native seeding
Restoration pathway T3A
State 3 to 2
Long-Term fire suppression and/or land abandonment
Transition T3B
State 3 to 4
Agricultural conversion via tillage, seeding, and non-selective herbicide
Transition R3A
State 3 to 5
Site preparation, tree planting, non-native species control and native seeding
Restoration pathway T4A
State 4 to 2
Long-term fire suppression and/or land abandonment
Restoration pathway T4B
State 4 to 3
Cultural treatments are implemented to increase forage quality and yield
Transition R4A
State 4 to 5
Site preparation, tree planting, non-native species control and native seeding
Restoration pathway T5A
State 5 to 2
Long-term fire suppression and/or land abandonment
Restoration pathway T5B
State 5 to 3
Cultural treatments are implemented to increase forage quality and yield
Restoration pathway T5C
State 5 to 4
Agricultural conversion via tillage, seeding and non-selective herbicides
