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 transition zone between the eastern deciduous forests and the tallgrass prairies. The heterogeneous topography of the area results in variable microclimates and fuel matrices that in turn are able to support prairies, savannas, woodlands, and forests. Loess Upland Prairies form an aspect of this vegetative continuum. This ecological site occurs on upland hillslopes and high stream terraces on moderately well to well-drained soils. Species characteristic of this ecological site consist of sun-loving, fire-adapted herbaceous vegetation.

Fire is a critical disturbance factor that maintains Loess Upland Prairies. Fire intensity typically consisted of periodic, low-intensity surface fires occurring every 1 to 3 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, driving large game, improving grazing and browsing habitat, agricultural clearing, and enhancing vital ethnobotanical plants (Barrett 1980).

Drought and herbivory by native ungulates have also played a role in shaping this ecological site. The periodic episodes of reduced soil moisture in conjunction with the moderately well to 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. Bison (Bos bison) grazing, while present, served a more limited role in community composition and structure than lands further west. Prairie elk (Cervus elaphus) and white-tailed deer (Odocoileus virginianus) likely contributed to woody species reduction but are also considered to be of a lesser impact compared to the west (LANDFIRE 2009). When coupled with fire, periods of drought and herbivory can further delay the establishment of woody vegetation (Pyne et al. 1996).

Today, Loess Upland Prairies are limited in their extent, having been type-converted to agricultural production land. Remnants that do exist show evidence of indirect anthropogenic influences from fire suppression and non-native species invasion. A return to the historic plant community may not be possible following extensive land modification, but long-term conservation agriculture or prairie 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 a mesic tallgrass community, dominated by herbaceous vegetation. The two community phases within the reference state are dependent on fire. The intensity and frequency alter species composition, cover, and extent, while regular fire intervals keep woody species from dominating. Drought and herbivory have more localized impacts in the reference phases, but do contribute to overall species composition, diversity, cover, and productivity.

Community Phase 1.1 Leadplant/Big Bluestem – Pinnate Prairie Coneflower – Sites in this reference community phase are dominated by a mix of grasses and forbs with patchy shrubs. Vegetative cover is continuous (95 to 100 percent) and plants can reach heights between 3 and 6 feet tall (LANDFIRE 2009; NatureServe 2015). Big bluestem, Indiangrass, little bluestem, sideoats grama, and prairie dropseed (Sporobolus heterolepis (A. Gray) A. Gray) are the dominant warm-season grasses present on the site. Characteristic forbs can include pinnate prairie coneflower, wild bergamot (Monarda fistulosa L.), and compassplant (Silphium laciniatum L.). Low shrubs, such as leadplant and prairie rose, can be sparsely present (NatureServe 2015). Fire with low intensity will maintain this community phase, but a hot, replacement fire will shift the site to community phase 1.2 (LANDFIRE 2009).

Pathway 1.1A – Hot, replacement fire every 1 to 3 years.

Community Phase 1.2 Big Bluestem – Annual Ragweed – This reference community phase represents a successional shift following a major disturbance event. A high intensity fire will remove or significantly reduce the shrub component as well as create open pockets of bare ground where annuals, biennials, and short-lived perennials can germinate. Perennial, warm-season grasses are still dominant on the site, but forb diversity will include opportunistic species such as annual ragweed (Ambrosia artemisiifolia L.) and clammy groundcherry (Physalis heterophylla Nees). Vegetative regrowth following the fire is rapid, and plants can reach maximum heights by the end of the growing season. A couple years without fire allows the community to mature, shifting back to phase 1.1 (LANDFIRE 2009).

Pathway 1.2A – Natural succession as a result of a brief fire-free period.

Transition 1A – Long-term fire suppression transitions the site to the fire-suppressed scrub state (2).

Transition 1B – Cultural treatments to enhance forage quality and yield transitions the site to the forage state (3).

Transition 1C – Tillage, seeding of agricultural crops, and non-selective herbicide transition the site to the cropland state (4).


STATE 2 – FIRE-SUPPRESSED SCRUB STATE

Long-term fire suppression can transition the reference tallgrass prairie community into a woody-invaded shrub-prairie state. This state is evidenced by a well-developed shrub layer and sparse trees (LANDFIRE 2009). Proximity to lands that have been altered provide opportunities for non-native invasive species to readily colonize this state, thereby reducing the native biodiversity and changing the vegetative community.

Community Phase 2.1 Roughleaf Dogwood – Multiflora Rose/Big Bluestem – Kentucky Bluegrass – This community phase represents the early stages of fire-suppression. In as little as six fire-free years, the prairie is disrupted and succeeded by woody shrubs. Native species – e.g., roughleaf dogwood (Cornus drummondii C.A. Mey), black cherry (Prunus serotina Ehrh.), black raspberry (Rubus occidentalis L.) – and non-native species – e.g., multiflora rose (Rosa multiflora Thunb.) – can form dense thickets with cover reaching up to 30 percent and plant heights as tall as 9 feet (LANDFIRE 2009). Some native prairie plants will persist, but non-native herbaceous species tolerant of shading encroach on the site.

Pathway 2.1A – Continued fire suppression in excess of 20 years.

Community Phase 2.2 Common Hackberry/Roughleaf Dogwood – Multiflora Rose/Kentucky Bluegrass – Sites falling into this community phase have a well-established shrub layer, and scattered trees begin to develop in the continued absence of fire. The shrub canopy can be diverse, including both native and non-native species. Roughleaf dogwood, black cherry, black raspberry, and eastern poison ivy (Toxicodendron radicans (L.) Kuntze) are common natives, and multiflora rose is a frequently invading non-native. Common hackberry (Celtis occidentalis L.), silver maple (Acer saccharinum L.), and elms (Ulmus L.) are the most common native trees present. The non-native white mulberry (Morus alba L.) can also be encountered.

Pathway 2.2A – Single large disturbance event such as selective removal of woody species.

Transition 2A – Cultural treatments to enhance forage quality and yield transitions the site to the forage state (3).

Transition 2B – Tillage, seeding of agricultural crops, and non-selective herbicide transition this site to the cropland state (4).

Restoration 2A – Site preparation, invasive species control, and seeding native species transition this site to the reconstructed tallgrass prairie state (5).


STATE 3 – FORAGE STATE

The forage state occurs when the site 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 (Smith 1998). 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 Phase 3.1 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).

Pathway 3.1A – Mechanical harvesting is replaced with domestic livestock utilizing continuous grazing.

Pathway 3.1B – Mechanical harvesting is replaced with domestic livestock utilizing rotational grazing.

Community Phase 3.2 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.

Pathway 3.2A – Domestic livestock are removed, and mechanical harvesting is implemented.

Pathway 3.2B – Rotational grazing replaces continuous grazing.

Community Phase 3.3 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 3.3A – Continuous grazing replaces rotational grazing.

Pathway 3.3B – Domestic livestock are removed, and mechanical harvesting is implemented.

Transition 3A – Land abandonment transitions the site to the fire-suppressed scrub state (2).

Transition 3B – Tillage, seeding of agricultural crops, and non-selective herbicide transition this site to the cropland state (4).

Restoration 3A – Site preparation, invasive species control, and seeding native species transition this site to the reconstructed tallgrass prairie state (5).


STATE 4 – CROPLAND STATE

The low topographic relief across the MLRA has resulted in nearly the entire area being converted to agriculture (Eilers and Roosa 1994). 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 Phase 4.1 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).

Pathway 4.1A – Tillage operations are greatly reduced, crop rotation occurs on a regular interval, and crop residue remains on the soil surface.

Pathway 4.1B – Tillage operations are greatly reduced or eliminated, crop rotation occurs on a regular interval, crop residue remains on the soil surface, and cover crops are planted following crop harvest.

Community Phase 4.2 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.

Pathway 4.2A – Intensive tillage is utilized, and monoculture row-cropping is established.

Pathway 4.2B – Cover crops are implemented to minimize soil erosion.

Community Phase 4.3 Conservation Tillage with Cover 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 4.3A – Cover crop practices are abandoned.

Pathway 4.3B – Intensive tillage is utilized, cover crops practices are abandoned, monoculture row-cropping is established, and crop rotation is reduced or eliminated.

Transition 4A – Land abandonment transitions the site to the fire-suppressed scrub state (2).

Transition 4B – Cultural treatments to enhance forage quality and yield transitions the site to the forage state (3).

Restoration 4A – Site preparation, invasive species control, and seeding native species transition this site to the reconstructed tallgrass prairie state (5).


STATE 5 – RECONSTRUCTED TALLGRASS PRAIRIE STATE

Prairie reconstructions have become an important tool for repairing natural ecological functions and providing habitat protection for numerous grassland dependent species. Because the historic plant and soil biota communities of the tallgrass prairie were highly diverse with complex interrelationships, historic prairie replication cannot be guaranteed on landscapes that have been so extensively manipulated for extended timeframes (Kardol and Wardle 2010; Fierer et al. 2013). 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 natural ranges of stress and disturbance, and create and maintain positive biotic and abiotic interactions (SER 2002). The reconstructed prairie state is the result of a long-term commitment involving a multi-step, adaptive management process. Diverse, species-rich seed mixes are important to utilize as they allow the site to undergo successional stages that exhibit changing composition and dominance over time (Smith et al. 2010). On-going management via prescribed fire and/or light grazing can help the site progress from an early successional community dominated by annuals and some weeds to a later seral stage composed of native, perennial grasses, forbs, and a few shrubs. Establishing a prescribed fire regimen that mimics natural disturbance patterns can increase native species cover and diversity while reducing cover of non-native forbs and grasses. Light grazing alone can help promote species richness, while grazing accompanied with fire can control the encroachment of woody vegetation (Brudvig et al. 2007).

Community Phase 5.1 Early Successional Reconstructed Tallgrass Prairie – This community phase represents the early community assembly from prairie reconstruction and is highly dependent on the seed mix utilized and the timing and priority of planting operations. The seed mix should look to include a diverse mix of cool-season and warm-season annual and perennial grasses and forbs typical of the reference state (e.g., big bluestem, Indiangrass, little bluestem, pinnate prairie coneflower). Cool-season annuals can help provide litter that promotes cool, moist soil conditions to the benefit of the other species in the seed mix. The first season following site preparation and seeding will typically result in annuals and other volunteer species forming a majority of the vegetative cover. Control of non-native species, particularly perennial species, is crucial at this point to ensure they do not establish before the native vegetation (Martin and Wilsey 2012). After the first season, native warm-season grasses should begin to become more prominent on the landscape.

Pathway 5.1A – Selective herbicides are used to control non-native species, and prescribed fire and/or light grazing helps to increase the native species diversity and control woody vegetation.

Community Phase 5.2 Late Successional Reconstructed Tallgrass Prairie – Appropriately timed disturbance regimes (e.g., prescribed fire) applied to the early successional community phase can help increase the beta diversity, pushing the site into a late successional community phase over time. While prairie communities are dominated by grasses, these species can suppress forb establishment and reduce overall diversity and ecological function (Martin and Wilsey 2006; Williams et al. 2007). Reducing accumulated plant litter from perennial bunchgrasses allows more light and nutrients to become available for forb recruitment, allowing greater ecosystem complexity (Wilsey 2008).

Pathway 5.2A – Reconstruction experiences a decrease in native species diversity from drought or improper timing of management actions (e.g., reduced fire frequency, use of non-selective herbicides).

Transition 5A – Land abandonment transitions the site to the fire-suppressed scrub state (2).

Transition 5B – Cultural treatments to enhance forage quality and yield transition the site to the forage state (3).

Transition 5C – Tillage, seeding of agricultural crops, and non-selective herbicide transition this site to the cropland state (4).