Tallgrass Prairie (Interior) - IA

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The tallgrass prairie includes range vegetation that can be visualized as consisting of two major forms: 1) once-vast grasslands in the continental interior of North America and 2) zones of grasslands and marshes along the shores of North America extending from the Atlantic Coast through the Gulf of Mexico. Climate of the former is continental; that of the latter is maritime. Species composition (including dominants) of plants and animals is similar (sometimes nearly the same) on both of these two basic forms or expressions of tallgrass range. Differences in species in the pre-Columbian ecosystems of these forms was probably not great as there were similar species (= similar ecological niches) or ecological equivalents among range types in them. For example, the dominant climax gallinaceous birds of the interior tallgrass bluestem prairies (eg. Flint Hills and Osage Questas of Kansas and Oklahoma), Gulf of Mexico coastal cordgrass prairies and marshes, and small patches of northern cordgrass prairies along the Atlanic Coast were one subspecies of the grouse known as greater prairie chicken (Tympanuchus cupido pinnatus), Attwater prairie chicken (T. cupido attwateri), and heath hen (T. cupido cupido), respectively. Of course the heath hen is extinct, the Attwater in dire trouble, and the greater is declining, but such similarity in taxa of the major grassland bird of each of these recognized generic grasslands illustrated the ecological relatedness of these major forms of tallgrass grasslands and various range cover types therein.

For pedagogic purposes and convenience (ease of viewing examples) the tallgrass prairie sub-formation (or sub-biome) of North Amereican grasslands was divided into two separate chapters designated as interior tallgrass prairie and coastal tallgrass prairie. There are marshes (herbaceous plant communities usually with the soil surface covered by standing water most of the year or growing season) in (or affilitated with) both of these major forms of tallgrass prairie. Marshes have often interpreded as different from grasslands, perhaps even as separate biomes. Natural distinction between prairie (dominant plants are species of grasses hence grassland) and marsh (dominant plants include species of grasslike plants such as sedges, bulrushes or tules, rushes, cattails, etc. as well as grasses). Designations and differences are often unclear or even arbitrary as, for example, distinction between wet prairie and marsh. The same dominant and associate species of plants are sometimes common to adjacent grassland and marsh. This condition is more common in coastal prairies and marshes than in interior prairies and associated wetlands. Generally marshes are more common and prominent plant communities in coastal than in interior grassland vegetation. Such was reflected in names of these two major forms of tallgrass prairie.

It could be argued that the term prairie could or should encompass both grassland and marsh thereby uniting these two general units of range vegetation. Perhaps prairie could be interpreted so as to include both grassland and marsh. Such a union is not possible given the traditional designation of vegetational or land resource area 2 in Texas as Coastal Prairies and Marshes. The long-standing convention of this "purple-pedigreed' title was retained in the current publication. Rangemen simply have to learn to live with a certain amount of ambiguity.

In the Gulf Prairies and Marshes there are range plant and animal communities that are not strictly speaking dominated by tallgrass species. These include such habitats as seashores and salt flats. Such range vegetation or plant and animal communities are, however, part of the Gulf Prairies and Marshes landscape or greater ecosystem (when seen from pserpective of Landscape Ecology or Ecosystem Ecology). In context of a publication devoted to range types these units of range vegetation were distinctive and different enough from described rangeland cover types (Shiflet, 1994)-- and obviously essential or integral to development and functioning of recognized range types-- that such vegetation was included herein.

The ultimate source for native vegetation of the tallgrass prairie (and true prairie) is North American Prairie (Weaver, 1954). Definitive reference for tallgrass prairie from an ecosystem perspective is that of Kucera (in Coupland, 1992, ps. 227--268).

Note on organization (and navigational directions): most of the range plant species listed for examples of tallgrass prairie range types presented in this chapter are in the sister chapter entitled, Tallgrass Prairie (Interior)- I B.

1. The Virgin Prairie- Tallgrass prairie; bluestem pastures. Physiography and vegetation of the Flint Hills. Tallgrass prairie dominated by "The Four Horsemen of the Prairies": big bluestem (Andropogon geraldii= A. furcatus), little bluestem (A. scoparius = Schizachyrium scoparium), Indiangrass (Sorgastrum nutans), and switchgrass (Panicum virgatum).Konza Prairie,

Riley County, Kansas. Estival aspect, July. FRES No. 39 (Prairie Ecosystem). K-66 (Bluestem Prairie). SRM 710 (Bluestem Prairie) or SRM 601 (Bluestem Prairie). "Tall-Grass" Series, 142.11, of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Flint Hills Ecoregion, 28a (Chapman et al., 2001).

* Note: In the North American biotic community classification of Brown et al. (1998, p. 40) all cover types within the tallgrass prairie were encompassed by the generic--and overly broad-- designation of Bluestem "Tall-Grass" Series, specifically Bluestem "Tall-Grass Series, 142.11 of Plains Grassland biotic community, 142.1, except for the Beardgrass Series of Gulf Coastal Grassland. Even the latter was too general for the less diverse Gulf Coastal Tallgrass Prairie. The title of Bluestem "Tall-Grass" Series was not repeated again because there are numerous and more specific rangeland cover types within the tallgrass prairie. The better known Kuchler unit, Bluestem Prairie, was shown throughout because it was basis of the Prairie Ecosystem of the long-standing Forest and Range Ecosystem series.

2. Tallgrass prairie transect- "Compressed" along this length (approximately 80 yards) of Flint Hills sod is the complete array of all major tallgrass prairie communities going from wet bottomland to shallow, rocky hilltop. In the immediate left foreground is an eastern gamagrass consociation (a zone of "pure" eastern gamagrass or natural single- species stand) except for two or three patches of canada or nohe third "strip", of two species: 1) big bluestem (far left patch of shorter grass) which is a short-shoot grass that does not elongate its culm until late summer so infloresences have just emerged and 2) a bottomland ecotype of switchgrass (far right midground that has fully emerged its panicle inflorescences that have a reddish cast). Switchgrass is a long-shoot tallgrass that elongates its culm and apical meristem relatively early in the growing season. The fourth zone is Indiangrass which is also a long-shoot tallgrass species of about the same height as the switchgrasss and, being behind it, is not visible from this vantage point. The hillside in the background is a "three-way" dominance mixture of big bluestem, Indiangrass, and sideoats grama farily evenly distributed but with the three respective species becoming progressively less dominant as they populate the hillside until sideoats grama forms a “pure” single-species stand at the summit except for scattered compassplants which are visible against the skyline. The main associate interspersed in the eastern gamagrass and prairie cordgrass is the prominent, tall, yellow composite, wholeleaf rosin-weed (Silphium integrifolium). The mid-height graminoid in the bottom of the draw in far right foreground is another Carex species.

This zonal distribution of species in the vegetation is due to soil moisture (and soil depth largely as a determinant of soil water). This is the phenomenon that F.E. Clements (1920, p. 26; Weaver and Clements, 1938, p. 203) explained by the term chresard ("amount for use") or soil water available for use by plants. Like most of Clements’ terms this one never caught on (perhaps fortunately so). It seems synonymous with the term and concept of field capacity water for mesophytes, but Clements used chresard as one major factor to explain distribution of plants. The zonation of tallgrass prairie species along this chresard gradient is gradation from most mesic (or maybe hydric) to least mesic or most xeric. It is a graphic reminder of this major fact of life on the range:

Available (=effective soil water is the most universally limiting factor in range plant survival, growth, and reproduction. Soil water available for plant use is the single most important factor in determining species distribution, plant community productivity and, ultimately, yield of range animal products (meaning ranch income).

"All Hell needs is water." (Attributed to General Phillip H. Sheridan as his comeback when he was told that all Texas needed was water. This assessment was a response to Sheridan’s initial conclusion: “If I owned Hell and Texas I would live in Hell and rent out Texas”.

Konza Prairie, Riley County, Kansas. Estival aspect, August. FRES No. 39 (Prairie Ecosystem), K-66 (Bluestem Prairie). SRM 601. Flint Hills Ecoregion, 28a (Chapman et al., 2001).

3.Osage landscape- Landscape scale views of the Osage Questas portion of the greater or general Flint Hills Region, all of which is in the Osage Plains section of the Central Lowlands physiographic province (Fenneman, 19ps. 605-618). Here in the southernmost extension of the Flint Hills this has traditionally been known as the Prairie Plains (Fenneman, 1938, ps. 616-617) parent material is alternatively limestone or sandstone. The rangeland presented here was primarily in the limestone or "strong grass" country of the Osage where herbage is generally more nutritious and cattle gains greater than on the sandstone-derived soils. Cross Timbers vegetation develops on sandstone-derived soils however.

This tallgrass prairie-Cross Timbers range was on the western edge of the Prairie Peninsula of Transeau (1935). The patches of woodland are part of the Eastern Cross Timbers dominated by post oak (Quercus stellata) and blackjack oak (Q. marlandica). Soils are the Stephenville-Darnell series that comprise Sandy Savanna to Shallow Savanna range sites. Tallgrass prairie vegetation is the classic Four Horsemen of the Prairies (big and little bluestems, Indiangrass, and Switchgrass) with wild alfalfa (Psoralea tenuifolia) as dominant forb This classic tallgrass prairie grassland was on Coweta and Bates soil series that made up Shallow Praiirie and Loam Prairie range sites.

Prairie Peninsula (Transeau, 1935) was treated immediately below.

Osage County, Oklahoma. May; vernal aspect. This natural vegetational mosaic was a combination of FRES No. 39 (Prairie Grasslands Ecosystem), K-66 (Bluestem prairie) and FRES No. 15 (Oak-Hickory Forest Ecosystem), 72 (Oak Savanna). SRM 710 (Bluestem Prairie) and SRM 731 (Cross Timbers- Oklahoma). "Tall-Grass" Series, 142.11, of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Flint Hills Ecoregion, 28a (Woods et al., 2005).

4. Woods in the prairie- Edges of tallgrass bluestem-dominated prairie and gallary forest in the Flint Hills. The prairie range was comprised primarily by the Four Horseman of the Prairies along with eastern gamagrass. Baldwin ironweed (center of first photograph) and whole-leaf rosinweed (Silphium integrifolium) (lower left corner of second photograph) were common forbs. Characteristically both forbs were composites.

Gallary or fringing forest has generally been applied to tropical vegetation as in regions of rain-forests and savannahs, but ecologists have adopted its use for temperate regions as well. In general a gallary forest is one that follows along streams from stream bank to exterior of the forest where outer edge of this forest contacts adjoining and different (often drastically so) vegetation such as grassland or savanna. Gallary forest includes the riparian zone but extends beyond it to include all woodland or forest that extended consistently from creek, river, etc. to another kind of vegetation. Gallary forests are by definition limited to stream-based, stream-contacting forests that meet another formation or some lower level of major plant community. Gallary forests are not to be confused with streamside forest vegetation bounded by forests (even if this forest plant community differs from the woody riparian vegetation).

In the terminology and basic concepts of Landscape Ecology, gallary forests are corridors within a matrix of non-forest vegetation or corridors that connect different landscape matrices and/or patches.Gallary forest were extremely valuable (especially given their limited size as corridors) native forest communities in pre-Columbian grassland and savannas, especially of central North America.

Most gallary and riparian forests were treated in this publication inside various chapters of Forests and Woodlands. This example of gallary forest was included here for consistency and comprehensiveness in describing tallgrass prairie range vegetation. Another example of gallary forest-tallgrass prairie edge (forest-grassland ecotone) was presented later in this Tallgrass Prairie (Interior) chapter in context of the North American Prairie Peninsula.

Major tree species of the gallery forest introduced here were given in the caption for the next succeeding photograph that revealed the interior of this fringing forest with its general structure.

Kings Creek, Konza Prairie, Riley County, Kansas. Estival aspect, early August. FRES No. 39 (Prairie Grasslands Ecosystem), K-66 (Bluestem Prairie) and FRES No. 15 (Oak-Hickory Forest Ecosystem), K-73 (Mosaic of Bluestem Prairie [K-66] and Oak-Hickory Forest [K-91]). SRM 710 (Bluestem Prairie) and SRM 731 (Cross Timbers- Oklahoma). "Tall-Grass" Series, 142.11, of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Flint Hills Ecoregion, 28a (Chapman et al., 2001).

5. Gallery forest in Flint Hills tallgrass prairie- This is a veiw standing in virgin tallgrass prairie of the Four Horsemen prairie grasses (big bluestem is dominant and eastern gamagrass is the main associate with Baldwin ironweed the conspicuous forb) looking into a gallery forest dominated by bur and chinquapin oaks (Quercus muhlenbergii) and common or western hackberry (Celtis occidentalis) with green ash (Fraxinus pennsylvanica), sycamore (Platanus occidentalis) and American or white elm (Ulmus americana) as associates. A gallery forest describes the narrow forest community that grows along the corridor of a water course. It is not only the riparian vegetation that grows directly in the hydric habitat but also forest vegetation growing beyond wet soil. The physiogonomy and mosaic pattern at this close distance resembles that of the Prairie Peninsula of Transeau (1935).

6. Patch of hardwoods in an Osage prairie- Interior of a gallary forest or, more specifically, an open forest or big tree-savanna by an ephemeral stream or drainage within (completely surrounded by) tallgrass prairie. This bottomland savanna or open forest consisted of black oak (Quercus velutina), bur oak, sycamore green ash, and red mulberry (Morus rubra) with a grassy understorey of tallgrass species including the obvious eastern gamagrass plus big bluestem, switchgrass, Indiangrass, beaked panicgrass (Panicum anceps), Canada wildrye, and purpletop (Tridens flavus) plus tickclover (Desmodium spp.) as the major herbaceous legume. Blackberry (Rubus spp.) was the dominant understorey shrub. It grew in colonies (ie. blackberry patches). Trumpet creeper (Campsis radicans) grew up most of the large trees thereby forming an interrupted layer of liana (woody vine). Poison oak (Rhus toxicodendron= Toxicodendron radicans= Rhus radicans) was also present as a liana though at less abundance.

In the conceptual framework of Landscape Ecology this open canopy bottomland hardwood forest or hardwood-tallgrass savanna (depending on interpretation) was a patch within a matrix of tallgrass prairie.

Structure and species composition of this bottomland savanna or open forest range was remarkably similar (almost identical by standards of vegetation) to the gallary forest. The bottomland savanna was in the southern Osage Questas portion of the Central Lowlands physiographic province whereas the gallary forest treated immediately above was in the Flint Hills portion of the Central Lowlands. Both of these samples of tree-dominated vegetation were quite similar to the ecotone between tallgrass prairie and oak-hickory forest in the Ozark Plateau to the east and the Cross Timbers to the west and south. In fact, the Western Cross Timbers extend through this same area so that distinction among these tree-dominated plant communities becomes problematic and of little practical importance.

Examples of this ecotonal or savanna range vegetation was treated in part at this juncture to show continuity within such natural vegetation and for comprehensive coverage of range plant communities found within the general tallgrass prairie region and the partially conterminous Prairie Peninsula that prior to modification by european man extended eastward and northward from southern and central parts of the tallgrass prairie. An example of actual or per se Prairie Peninsula range vegetation was included later in this chapter.

Chapman-Barnard Ranch, Osage County, Oklahoma. Vernal aspect. May. General range vegetation was FRES No, 39 (Prairie Ecosystem). K-66 (Bluestem Prairie). SRM 710 (Bluestem Prairie). Specific vegetation of this bottomland hardwood-tallgrass savanna was FRES NO. 15 (Oak-Hickory Forest Ecosystem), K-73 (Mosaic of Bluestem Prairie [K-66] and oak-hickory [K-91]), closest SRM designation was SRM 731 (Cross Timbers, Oklahoma) in Flint Hills Ecoregion, 28a (Woods et al., 2005).

Flint Hills form of tallgrass prairie- The Flint Hills is a specific portion of the Central Lowlands physiographic province. In common (informal or layman's usage) "Flint Hills" is used generically in reference to tallgrass prairie in Kansas and Oklahoma underlaid with limestone and sandstone rock which has protected the wonderful grasslands, some of the finest natural pasture on Earth, from the ravishes of the plow. In precise usage, Flint Hills is a much more restricted, physiographically defined geologic area. The Flint Hills, along with the Chautauqua Hills and Osage Cuestas or Questas are in the Osage Section of the Central Lowland physiographic province. Fenneman (1938, ps. 605-620 and, especially, ps. 614-616) remains the definitive authority. This is sometimes referred to as the Osage Plains as, for instance, by Orme (2002, ps. 343-345) which is another outstanding source.

The most accurate mapping of these various physiographic units may well be the Kansas Offficial Transportation Map (Kansas Department of Transportation, various years) which showed delinations within Kansas counties. This and the Kansas Geological Survey on the web (under GeoKansas) were obviously taken from the seminal work of Schoewe (1949).

The Flint Hills are regarded as a cuesta (Fenneman, 1938, ps. 606-609), a term of varied though similar meanings. American Geological Institute (Gary et al., 1972) offered two sikghtly different definitions of cuesta with the first being the one that most fit the usage applied to Flint Hills and Osage Cuestas: "A hill or ridge with a gentle slope on one side and a steep slope on the other; specifically, an asymmetric ridege (as in SW U.S.) with one face (dip slope) long and gentle and conforming with the dip on the resistant bed or beds that form it, and the opposite face (scarp slope) steep or even cliff-like and formed by the outcrop of the reisitant rocks, the formation of the ridge being controlled by the differential erosion of the genctly inclined strata."

The less resistant stratum (layer) of he Flint Hills was the

7. Framed in "Flint"- Landscape view of the picturesque and productive Flint Hills of Kansas, one of the largest remaining expanses of tallgrass prairie and, as often claimed, "God's own steer country". Grassland vegetation in this view was typical Four Horsemen of the Prairies grasses (big bluestem, little bluestem, Indiangrass, and swithcgrass).

There were at least two range sites on this tallgrass prairie landscape: 1) Loamy Upland in the foreground and 2) Limy Upland in background. Shallow Limy range site was behind the Limy Upland and not visible in this photograph. On the Loamy Upland range site big bluestem was the dominant and switchgrass of upland ecotype(s) was the associate. On the Limy Upland little bluestem was dominant while Indiangrass was the associate species. Sideoats grama dominated the Limy Upland range site with little bluestem being the associate. Rangeland viewed here constituted a toposequence, specifically a toposequence of soils or a catena. Catena was defined by the Soil Science Society of America (2001) as "a sequence of soils of about the same age, derived from similar parent material, and occurrring under similar climatic conditions, but having different characteristics due to variation in relief and in drainage". Soils constituted the primary basis of the range sites.

Geologically speaking the geomorphic/physiographic material of the Flint Hills is that of limestone, chert, and, even, some shale in bedrock tracing to the Permian Age. There is a "protecting mantle of flints" in some areas of the Flint Hills section of the Central Lowland province that are remains of a former peneplain. Geologic erosion of this flint left the deeper limestones and cherts so that the Flint Hills is a cuesta with distinct terraces and incised stream valleys that dissect this escarpment (Fenneman, 1938, ps. 606-609, 614-618).As used here "flint" was in reference to chert which is less water soluble than limestone so that with weathering and geologic erosion there remained remnants of chert ("flint" in this application) as gravel. This cherty gravel is the remains of Permian Age beds on the land surface that was once overlaid by inland seas (GeoKansas, Kansas Geological Survey, Undated). In other words, the Flint Hills are actually limestone and chert not flint at all.

To add further confusion, the designation of Flint Hills is misapplied or, at least, loosely if not carelessly applied as a generic or catchall category for all tallgrass prairie in Kansas and northern Oklahoma. In this loose usage "Flint Hills" includes the adjoining Smoky Hills to the west, adjoining Osage Cuestas to the east, the Chautauqua Hills to the southeast, and even the Great Bend and Arkansas River lowlands to the west. Similarity of vegetation across the natural grasslands of this region attest to the "veto power" of climate over edaphic and topographic features in final determination of regional climax vegetation. The Chautauqua Hills that were formed from Douglas sandstone (Fenneman, 1939, ps. 613-614) are the greatest deviation from the general tallgrass prairie vegetation. In this physiographic section the climax plant community is that of the Cross Timbers savanna rather than grassland communities.

Distinction and precise application of Flint Hills was followed herein as were later discussions of the range vegetation of adjoining physiographic sections of the Central Lowlands province. Proper designations as to physiographic and geologic bases of tallgrass prairie vegetation was of practical, utilitarian importance. For example, cattlemen have long known that performance of cattle, especially of stockers, is noticably greater for the "strong grass" on limestone-derived soils such as the Flint Hills in contrast to the "weaker grass" on soils that developed from sandstone like those of the Chautauqua Hills.

Marion County, Kansas. Mid-July (early estival aspect). FRES No. 39 (Prairie Ecosystem). K-66 (Bluestem Prairie). SRM 710 (Bluestem Prairie), perhaps more specifically, or SRM 601 (Bluestem Prairie ) generally. "Tall-Grass" Series, 142.11, of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Flint Hills Ecoregion, 28a (Chapman et al., 2001).

8. Blessed are the rocks- Flint Hills tallgrass prairie saved from the plow by an outcropping of limestone rocks from the Permian Period. This sequencial two-slide study of virgin sod in climax state was a rock outcrop form of the Limy Upland range site. This rangeland was shown in the background of the slide immediately above. Major grasses were little bluestem, big bluestem, Indiangrass, sideoats grama, and switchgrass (in that approximate order based on apparent foliar cover, plant density, and general abundance). Hairy grama and even buffalograss were locally abundant on shallow microsites, often beside outcropped rocks. Shrubs were not present and forbs were the same as absent.

This range had historically been summer steer range, but it had not been grazed during the current growing season. Big bluestem and Indiangrass had not begun to bolt (send up) sexual shoots in this immediate area. These same tallgrasses were in full-bloom stage on this same day about thirty miles to the south..

Note on equipment: this was the same slide (Fujichrome Provia 100F in this instance) scanned on two consecutive days with the same Epson scanner (Perfection 700) that had not been turned off, re-set, or had any other modification. This is simply an example of the inconsistency (lack of precision operation) of this apparatus. Never buy an Epson product.

9. Blessed are wet soils- Tallgrass prairie on a generally wet upland soil potentially dominated by switchgrass with big bluestem and, probably, prairie cordgrass as associates comprising the climax range vegetation. This stocker (steer) range was in Good range condition class with more big bluestem, tall dropseed (Sporobolus asper var. asper), Indiangrass, and sideoats grama relative to switchgrass than would be expected on Excellent condition range for this Clay Upland range site. There was relatively little prairie cordgrass and no visible plants of eastern gamagrass, two species that dominated more mesic forms of Clay Upland elsewhere in the Flint Hills (see below). The potential (climax) species composition of this grassland community was not known precisely, but it was most probably a switchgrass-prairie cordgrass form of wet tallgrass prairie though still with plenty of big bluestem.

The major forbs on this steer pasture were Baldwin ironweed (Vernonia baldwinii var. interior), wooly vervain (Verbena stricta), and wild alfalfa or manyflowered scurfpea (Psoralea tenuifolia var. floribunda). There were no shrubs on this prairie range. Annual burning is widespread in this locality and this pasture undoubtedly had been burnt countless times by both redman and whites.

Poor drainage can, like stoniness, be an edaphic feature that precludes tillage. Most field crops are mesophytes that require at least moderately well-drained soils. Poorly drained soils, especially thost that remain near the saturation point for prolonged periods, are unfit or, at least, marginal for production of row crops and small grains. Such wet soils are farmable only with expensive drainage. If in addition to excessive wetness such soils are not overly fertile to begin with and/or are in areas prone to drought it is not economically feasible to try to farm them. Instead, the native wet grassland is left for use as range. This is why rangemen sing praises for poorly drained as well as rock-strewn soils.

10. Famed Flint Hills-Strong grass = good steer country. Four Horsemen grasses with lots of forbs such as lead plant (Amorpha canescens) and wild alfalfa or slender scurfpea (Psoralea tenuifolia).

Riley County, Kansas. July.FRES No. 39 (Prairie Ecosystem). K-66 (Bluestem Prairie). SRM 710 (Bluestem Prairie), perhaps more specifically, or SRM 601 (Bluestem Prairie ) generally. "Tall-Grass" Series, 142.11, of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Flint Hills Ecoregion, 28a (Chapman et al., 2001).

11. Flint Hills bluestem prairie range- Long-yearling (more like just shy of two-year-olds) steers (some weighing in excess of 700 pounds) grazing tallgrass prairie in the Flint Hills cuesta (so named for a thin, veneerlike mantle of flint and chert overlaying Permian limestone). This is an example of the famed Flint Hills of Kansas (the formation extends some distance into northern Oklahoma) often regarded as one of the greatest natural cattle pastures on Earth and second only to the equally famous and fabulous tallgrass prairie of the Sand Hills of Nebraska.

Precisely speaking, much of the general tallgrass prairie pasture region in central and eastern parts of Oklahoma, Kansas, and Nebraska and the western portion of Missouri that are known generically as the Flint Hills are actually several geologically distinctive portions of the Osage Section of the Central Lowland physiographic province. Even in Kansas, most of the tallgrass (bluestem) prairie was in geologic units separate and distinct from the Flint Hills as strictly defined geologically. These other physiographic units include the Cherokee Prairie, Osage Questa, Chautauqua Hills, Glaciated Region, and Wellington and McPherson Lowlands. The Osage Questa and Flint Hills units with their limestone excarpments and adjacent proximity closely resemble each other (as can be seen by comparison of slides from both). The Chautauqua Hills sub-province is distinguished geologically by thick sandstone bedrock, but as it wedges up into the Osage Questa as a narrow peninsula and forms part of historical political units and famous grazing grounds like Osage County (Reservation), Oklahoma this distinctive unit is lumped as "Flint-- sometimes, Osage-- Hills".

Students are referred throughout this web publication to the timeless classic two-volume set on United States physiography by Fenneman. For geologic units just listed see Fenneman (1938, ps. 605-618). In some years the Kansas Official Transportation Map clearly delinates these physiographic units.

Big bluestem was easily the dominant on this steer range with Indiagrass and upland switchgrass coming in as "runners-up". Prairie dropseed (Sporobolus heterolepis), little bluestem, hairy grama (Bouteloua hirsuta), sideoats grama (B. curtipendula), and buffalograss (Buchloe dactyloides) were well-represtented but relatively rare. The most common forb was Illinhois bundleflower, but it had been grazed so heavily as to be less conspicuous than invader forbs like Baldwin ironweed and western yarrow (Achillea millefolium). A fenceline contrast comparing utilization of palatable forbs under heavier early stocking (Intensive Early Stocking or a grazing practice similar to it) for this same pasture and herd was presented in the following slide.

Butler County, Kansas. Early estival aspect, June. Viewers should bear in mind that big bluestem, the dominant species of the vegetation seen here, is a short shoot or "culmless" grass that does not elongate the culm above the basal leaves until late in the growing season (usually late July or early August in this location). As such, on bluestem prairie the namesake "tallgrass" growth habit does not occur (and therefore the characteristic physiogonomy of tallgrass prairie is not apparent) until late in the growing season. This is often in the autumnal aspect.

FRES No. 39 (Prairie Grassland Ecosystem), K-66 (Bluestem Prairie). SRM 601 (Bluestem Prairie), generally, or SRM 710 (Bluestem Prairie), more specifically. "Tall-Grass" Series, 142.11, of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Flint Hills Ecoregion, 28a (Chapman et al., 2001).

12. Visual comparison of ungrazed vegetation of bluestem prairie to that grazed by steers under heavier stocking early in the growing season- The ungrazed sward in the foreground was just outside the fence from the big bluestem-dominated range shown in the preceding slide. The area (both sides of the fence) had been burned off at onset of the current growing season (ie. "this spring"). Foliar cover of Illinois bundleflower was obviously greater where protected from grazing (which in this instance was more intensive utilization in the first part of the growing season). The grass in the immediate foreground was switchgrass, a long shoot species (one that elongates its culm and apical meristem higher earlier in the growing season).

Some cattlemen and range-leasing landowners either suspect or have concluded from their personal observations that heavier stocking, and consequent heavier degrees of use, early in the growing (= warm) season reduced populations of the more palatalble forbs, espcially legumes. Limited scientific data have been presented on responses of tallgrass prairie forbs to Intensive-Early Stocking. Owensby et al. (1988) reported that there were no consistent differences among stocking rate treatments on Intensive-Early Stocking of Flint Hills bluestem range, but their data and conclusions were of forb biomass only. Owensby et al. (1988) were not specific as to responses of individual species of forbs. It was not shown whether biomass, cover, density, etc. changed among decreasers like Illinois bundleflower and Maxmillian sunflower versus invaders like western ragweed. Was biomass of certain species the same under Intensive-Early Stocking as under traditional management or did biomass of decreasers decline while biomass of invaders increased (or vice versa)? The author was told by some landowners that management involving heavier stocking early in the growing season (either Intensive-Early Stocking or similar grazing management) had adverse impacts on decreaser forbs, especially legumes (personal communication, R.E. Lenington, DVM, MS, Cedar Vale, Kansas).

Flint Hills portion of the Osage Section of the Central Lowland physiographic province. Butler County, Kansas. Early estival aspect, June. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 601 (Bluestem Prairie), more generally, or SRM 710 (Bluestem Prairie), more specifically. "Tall-Grass" Series, 142.11, of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Flint Hills Ecoregion, 28a (Chapman et al., 2001).

13. Steer range in the famed Flint Hills- Tallgrass prairie range in the Osage Plains or Osage Section (the generic Flint Hills Region) has, under grazing use by whiteman, been steer rather than cow-calf country. The nutritive value and palatability of the tallgrass species coupled with gently rolling, easy traveling topography make for natural pasture that is ideal for putting on a lot of rapid, efficient gains (Average Daily Gain) on stocker cattle.

This big bluestem-dominated range was an example of the high-quality feed typical of this range type in central Kansas and northern Oklahoma. This is some of the fbest natural pasture for growing cattle available anywhere, period. On this range upland switchgrass and prairie dropseed were the two main associates to big bluestem. Other major grasses included little bluestem, Indiangrass, and Canada wildrye. Most of the forbs on this range were legumes including leadplant (Amorpha canadensis), both purple and white prairie clover (Petalostmon purpureum, P. candidum), Illinois bundleflower (Desmanthus illinoensis), and wild alfalfa (Psoralea tenuifolia). Tall gayfeather (Liatris aspera) was the most common, abundant composite. The main woody species was New Jersey tea (Ceanothus americanus).

The sward of this particular pasture was presented in the next two photographs.

Butler County, Kansas. Late June; early estival aspect. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 601 (Bluestem Prairie), more generally, or SRM 710 (Bluestem Prairie), more specifically. "Tall-Grass" Series, 142.11, of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Flint Hills Ecoregion, 28a (Chapman et al., 2001).

14. God's own pasture mix- Sward of the Flint Hills range introduced in the preceding two photographs. Main three grasses (in order) were big bluestem, upland switchgrass, and prairie dropseed. Little bluestem, Indiangrass, and Canada wildrye were next in oredr of abundance and overall importance. Most forbs were legumes including (in relative order) leadplant, purple prairie clover, white prairie clover, Illinois bundleflower, and wild alfalfa or slender scurfpea. Tall gayfeather was the most common composite. New Jersey tea was about the only woody species present.

15. Garden of Eden in the heartland- A south slope aspect of a hill in the Kansas Flint Hills in early summer presented in a series of wide angle (28mm) photographs to give something of a panaramic view of "God's own grassland". The dominant species was big bluestem and the associate species was switchgrass (an upland ecotype). This particular pasture had been a steer (spring-summer stockers) range for decades, but for whatever reason it had not been grazed by cattle this year (in the current warm-growing season). It was also routinely burnt as it had been in spring of the current year.

In the immediately succeeding slide this same pasture was shown in a previous year during which it had been grazed by steers which had been shipped just a few days prior to that photograph. In absence of cattle grazing this current year, leguminous prairie forbs had grown to large sizes (there had been minimal grazing by white-tailed deer on these legumes) so that they where very conspicuous. This included roundhead lespedeza (Lespedeza capitata), wild alfalfa or slim scurfpea (Psoralea tenuifolia), and lead plant (Amorpha canescenes). Other climax prairie forbs included tall goldenrod (Solidago altissima) and the highly palatable Maximillian sunflower (Helianthus-maximiliani). There were also some increaser forbs such as the perennial composite, Baldwin's ironweed (Veronia baldwinii).

There was some cover of the shrub, roughleaf or Drummond's dogwood (Cornus drummondii), but not nearly as much as in the adjoining, unburned, outer fencerow (see slides two slide-caption sets below the next landscape slide).

16. Flint Hills upland range- A predominately south slope on a large hill in the famed Flint Hills. Big bluestem was the dominant and switchgrass was the associate species. This range had been stocked with steers which had been shipped a few days prior to photograph.

Two "photo-quadrants" of this range vegetation just outside (across the fence from) this range were shown in the next two slides.

17. Burnt and unburnt- Fenceline contrast of the tallgrass prairie stocker range shown in the two immediately preceding two slide-caption sets. To left of the barbed wire fence was a tallgrass prairie range that had served as spring-summer steer range and that had been burnt almost yearly for decades. To right of the fence was the outer (outside) fencerow that had not been burned as frequently and not for several years prior to time of this slide.

The far greater cover of forbs and a few shrubs in the unburned side of the fencerow was glaringly obvious in this view. Dominant species on both sides of the fence were typical Four Horsemen of the Prairie grasses (big bluestem, little bluestem, Indiangrass, and switchgrsss in that order of cover/dominance), but there were more species and greater cover of forbs on the less frequently burned outer fencerow. These more abundant (greater cover and density) forbs visible in this slide included willowleaf sunflower (Helainthus salicifolius), Baldwin ironweed, and butterfly milkweed (Asclepias tuberosa). Also of greater cover and general abundance in the unburned side of the fence was that of the highly palatable shrub, New Jersey tea (Ceanothus americanus).

18. In the absence of periodic fire- Two closer-in "photoquadrants" of the less frequently burned vegetation of Flint Hills tallgrass prairie on the outside of the fenceline contrast presented in the immediately preceding slide-photo caption set. Grassland species in both slides included New Jersy tea, butterfly milkweed, willowleaf sunflower, Baldwin ironweed, American feverfew (Parthenium integrifolium), lead plant and big bluestem (the dominant grass species in the adjoining fired stocker range).

The same range plants were in both slides with differences in appearance being such details as flowering in butterfly milkweed in the second and not in the first "photoplot", more Baldwin ironweed cover and density in the second "photoplot", and greater cover of New Jersey tea in the first "photoplot".

19. Sward of upland tallgrass prairie- Two "photo-quadrants" of the south slope big bluestem-switchgrass Flint Hills range introduced in the four preceding slide-caption sets. These "photographic samples" were taken just a few yards across the fence from the south slope range where protection from grazing by steers permitted more detailed presentation of the species composition and physiogonomy of this climax range vegetation in early summer of a typical warm-growing season.

Herbaceous species present included big bluestem (the dominant), upland switchgrass (the associate), little bluestem, silver bluestem (Andropogon saccharoides), purple prairie clover, compassplant (Silphium laciniatum) willowleaf sunflower (Helianthus salicifolius), and tall gayfeather. The only shrub was New Jersey tea.

20. Burnt per nar every year- A bottomland habitat of Flint Hills tallgrass prairie dominated by the Four Horsemen tallgrass species in relative order of cover: switchgrass, big bluestem, Indiangrass, and little bluestem. There was some eastern gamagrass, tall dropseed, prairie dropseed, sideoats grama, silver bluestem, hairy grama, prairie cordgrass, Canada or nodding wildrye, buffalograss, etc. (the typical array of grass species). There were also a high density and cover of decreaser prairie forbs including Miximillian sunflower, purple prairie clover, and Mexican hat or prairie cone flower, but shrubs were not to be found.

The high burnline on the adult eastern cottonwoods (Populus deltoides subsp. deltoides) in the distance spoke to the frequent recurrent of prairie fire (prescribed burning). In fact, this stocker (steer) range was fired about every spring. This spring cattle had not yet been turned in on this pasture that was obviously a fine representative of the climax tallgrass prairie (Excellent range condition class).

21. Clean from annual burning- Two "photoquadrants" of the sward of the yearly burned stocker range of tallgrass prairie shown in the immediately preceding slide. The first "photoplot" (first slide) showed the vegetation of this range at farther distance while the second "photoplot" (second slide) presented the sward at closer distance.

Cattle had not been turned in on this range yet this spring which prompted the author to suspect that it was going to be used for prairie hay. It would certainly be the highest-quality prairie hay imaginable from the standpoint of species composition.

Switchgrass, big bluestem, Indiangrass, and little bluestem (in that relative order) comprised the vast "bulk" of the biomass in these two views of this climax range plant community. Silver bluestem, sideoats grama, tall dropseed, prairie dropseed, hairy grama, buffalograss Junegrass, prairie cordgrass, and Canada or nodding wildrye were also important grass species.

Major forb species included Maximillian sunflower, Mexican hat or prairie coneflower, purple prairie clover, leadplant, and slender or slim scurfpea. Note that most of the forbs in this climax range vegetation were composites and legumes.

With yearly burning nearly every spring, shrubs were about as "scarse as hen's teeth". The major, larger, herbivorous wildlife species was white-tailed deer (Odocoileus virginianus).

22. The dominant began to do its thing- Flint Hills tallgrass prairie at early bloom stage in big bluestem, the dominant species of this range and the Flint Hills section in general. Other major grasses on this particuar pasture included switchgrass, Indiangrass, little bluestem, prairie cordgrass, Canada wildrye, tall dropseed, and sideoats grama. Other grassses of infrequent occurrence ranged from eastern gamagrass to blue grama, hairy grama, and buffalograss. Almost all of these grasses were warm-season species except Canada wildrye which was about the only cool-season species that had any appreciable cover. The most common forb was Maximillian sunflower which was downright inconspicuous compared to its domination of the "prairie spotlight" when it reaches its maximum size and is in full bloom.

The first slide was of range vegetation in the pasture that was being grazed by steers. The second slide was outside the pasture with barbed wire readily visible. This second photograph presented big bluestem with zero livestock grazing (and no defoliation by deer being evident). Comparsion of ungrazed big bluestem (second slide) with big bluestem, as well as other tallgrass species, on the cattle-stocked pasture (first slide) served as a pictorial guide to light degree of use under light or conservative stocking.

This range was obviously pristine. Range condition class was Excellent and, to reiterate, degree of use was light. Stocking rate could have higher (greater degree of use such as, say, moderate) and still maintained the climax condition of this big bluestem-dominated range, but if a drought comes this range and the steers on it will come through in much better shape than those on which stocking rates were greater. The author "cherry picked" this example to show students the standard of perfection for native bluestem pasture in the Flint Hills being managed under tender, loving care of a faithful steward who knew (and cherished) what he had. Salute and three cheers!

23. Another grassland community (and a comparison of grazing intensity) in the Flint Hills- In contrast to the more typical big bluestem-dominated tallgrass prairie (the regional or zonal climax) that was presented immediately above, a more favorable (and more restricted) form of tallgrass grassland in the Central Lowlands province was this one seen here on which the potential (climax) dominant species was eastern gamagrass with prairie cordgrass being the associate species on moister habitats and big bluestem the associate on less mesic microsites. The surface of this rangeland had many local depressions that caught and retained greater quantities of water. Such microtopography provided more mesic microenvironments that supported these more moisture-requiring tallgrass species. This climax range vegetation was presented in the first of these two slides. It was on highway right of way immediately adjacent to (just outside the pasture fence) of an overgrazed pasture stocked with Angus cow-calf pairs.

The second slide was inside the overgrazed pasture which had been degraded through years of overuse. The overgrazed range was dominated by big bluestem on some local, less mesic habitats, but by hairy fimbry (Fimbristylis puberula) on the more mesic microenvironments. Big bluestem had more or less held its own on the drier habitats at some locations, but under continued overuse eastern gamagrass and prairie cordgrass had been grazed out and their places taken over by hairy fimbry. This was a textbook example of range deterioration through the process of retrogression, the retrograde (opposite the forward direction of plant sucession) changes in plant communities on the sere of this range site. Retrogrsssion occurred by the phenomenon of grazing selectivity through which range animals exhibit their preferences for species that are more palatable (have greater or higher palatability) to them.

Eastern gamagrass is one of the most palatable of all tallgrass species to bovids (cattle and buffalo). Prairie cordgrass is considerably less palatable to the bovids, but it is readily eaten by them and also subject to overuse. Hairy fimbry has a palatability somewhere between bailing wire and the box it came in. Any range consumer--other than rust (the kind left over after iron is consumed)--would prefer these two tallgrass species to hairy fimbry. Overstocking, too high a stocking rate, (with cattle on this range) led to overuse--excessive defoliation (exceeding proper degree of use)--of eastern gamagrass and prairie cordgrass. Overuse eventually weakened plants of these two palatable tallgrass species putting them at a competitive disadvantage to hairy fimbry which was not even touched (or same as) by cattle (or anything else with teeth). Over years or decades of overuse eastern gamagrass and prairie cordgrass were grazed out and replaced by hairy fimbry. At this point the condition of overgrazing had been reached. Longterm overuse leades to overgrazing. This change in species composition whereby more palatable species and those of higher successional order (climax decreaser species in this instance) were replaced by a plant species of lower successional order (hairy fimbry is a seral species that is an invader on this range site) was range deterioration Again, range degradation took place over time through the process of range retrogression as a function of grazing selectivity.

Degree of use (a function of stocking rate) on big bluestem had not been so great as to remove this decreaser from this range. In this example, it was only or, at least, primarily eastern gamagrass and prairie cordgrass that had been eliminated or greatly reduced by overgrazing. Important: successional status (decreaser, increaser, invader) is range site-specific. The range site of the pasture shown here was Clay Upland versus examples of Loamy Upland and Limy Upland range sites that were shown above. On Clay Upland there was proportionately greater soil moisture so that eastern gamagrass was the dominant and prairie cordgrass was the associate on low-lying wet areas and big bluestem was the principal associate on higher, drier habitats. On the still yet drier (less mesic) Loamy Upland range site big bluestem was the potential natural dominant. On Clay Upland, even if big bluestem (instead of hairy fimbry) dominated habitats once dominated by eastern gamagrass and/or prairie cordgrass the range would still be overgrazed. It would still be degraded because the climax dominants were replaced. Loss of these dominants was the only requirement, the only dynamic needed, in regards to the range vegetation for the condition of overgrazing to have occurred (or continue to be occurring). What plant species replaced the natural dominants was relevant only to the extent that replacement species were diagnostic as to severity of overgrazing (of range retrogression) and indicative (ie. such plants were indicator species) of how much time would be necessary for range recovery (or if range restoration was possible in practical time frames without corrective human action). To reiterate the point, even dominance by big bluestem, the major dominant of the regional climax, on a range site where some other plant species was (were) the natural dominant(s) would not override the successional fact of overgrazing. Range condition class and, more importantly, range trend is range site-specific. Successional status of range plant species and which plant species serve as indicator species varies as to range site.

Range sites have been regarded as the smallest, most distinctive unit of range having a characteristic vegetation that is practical to recognize, describe, map, etc. Out of practical necessity all mapping and descriptive units are generalities (though at a higher level of resolution) and products of compromises among defining vriables. There are variations and exceptions within all range sites the same as within soil series or other map units. All Clay Upland range sites share common specifics, but not all Clay Upland sites are the same. Microtopography with numerous depressions in the land surface on this rangeland made this form of Clay Upland distinct from others without such microrelief. The numerous "mini-ponds" of topographic "pockmarks" of this form of Clay Upland supported eastern gamagrass whereas switchgrass was the most mesic dominant on other Clay Upland sites in this area of Flint Hills that did not have as as large or as many depressions in the land surface.

Clarification: some range plant species are regarded as ice cream species: "An exceptionally palatable species sought and grazed frequently by livestock or game animals. Such species are often overutilized under proper grazing" (Jacoby, 1989). On some ranges and, more specifically, range sites eastern gamagrass is an ice cream species. It was NOT on the range (range site) presented. Instead, eastern gamagrass was the climax dominant over much of the Clay Upland especially in local depressions that were common on this range. Certainly, prairie cordgrass was not an ice cream species, least of all on Clay Upland. Replacement of these palatable species by hairy fimbry--that apparently nothing eats (or eats much of)--could not be written off as anything other than overgrazing. This dramatic change in dominance was not the phenomenon of ice cream species.

24. Could a'been a century ago- Texas longhorn cows and calves grazing the standard Four Horsemen (big bluestem, little bluestem, Indiangrass, and switchgrass) tallgrass prairie in the Flint Hills of the Osage Plains of the Central Lowlands physiographic province. Second slide stressed importance of abundant, clean water to livestock production and as a major tool in achieving Proper Distribution of Grazing Use, one of the Four Cardinal Principles of Range Manageament.

The tallgrass prairie shown here was roughly equal parts of the Four Horsemen with big bluesttem first among equals. Abundant cover of numerous other grasses ranging from prairie dropseed down to buffalograss (Buchloe dactyloides). Major forbs were Illinois bundleflower (Desmanthus illinoensis) and leadplant (Amorpha canadensis). About the only shrubs were buckbrush or coralberry (Symphoricarpos orbiculatus) and roughleaf or rough-leaved dogwood (Cornus drummondii). The only tree species present was bois d'arc, Osage orange, or hedgeapple (Maclura pomifera), a species probably introduced by whiteman (for a living fence; now used for the most durable of wood posts) and browsed by his cattle as evidenced by hedging of specimens in both of these photographs. Invasion by bois d'arc was not a good sign: if this alien (to these parts) brush species is not controlled it will take over this otherwise Excellent condition class range.

Crowley County, Kansas. Late June; late vernal asepct. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 601 (Bluestem Prairie), generally, or, probably more specifically, SRM 710 (Bluestem Prairie). Bluestem "Tall-Grass" Series, 142.11, of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Flint Hills Ecoregion, 28a (Chapman et al., 2001).

25. The way it looks when grazed by pairs- Appearance of big bluestem-little bluestem-Indiangrass-switchgrass-prairie dropseed range in the Flint Hills form of tallgrass prairie when grazed by cows and calves. Bluestem ranges grazed by stocker cattle (usually steers) consist almost entirely of current season's plant growth. There is little, if any, dead (standing or downed) herbage, aboveground plant material, on stocker pasture. This is because 1) grazing has been heavy enough and started early in the growing season that herbage from the previous growing season has been consumed and/or 2) ranges were burnt off prior to the warm growing season (fired in late winter or early spring) to increase availability to fresh feed.

By contrast, bluestem pastures used as range for cows and calves must of necessity be grazed lighter during the plant growing season in order to allow adequate accumulations of dead, dried herbage for winter feed for dry, pregnant, spring-calving cows or feed for wet cows and their calves if fall-calving is practiced. Also, ranges used for pasturing pairs are burnt less frequently so as to used herbage for feed instead of fuel.

The typical negative price structure for stockers (younger, smaller, lighter calves fetch more per cwt. than older, larger, heavier calves) means that profit can be made from stockers only by putting on sufficent weight during the grazing season (high Average Daily Gain; greater total weight gain over pasturing period). In other words, high levels of performance by individual animals--and range feed conditions conducive to high performance--is more critical for stocker than for cow-calf production. A corollary to this is that steers are run on ranges that provide more dry matter and more palatable, higher-quality (greater concerntrations of nutrients) cattle diets whereas cows and calves are grazed on ranges that furnish lower-quality, less palatable forage. Also pairs are stocked on rougher country: steep, rocky, brushy, etc. grazing lands do not provide overall good-quality pasturage that permits high levels of individual animal performance that are necessary for the typically low profit margins characteristic of stocker cattle.

Compare the appearance of this sward on a cow-calf range to that from steer ranges shown above.

Osage Cuestas (=Questas) form of tallgrass prairie- Presented in the following section were examples of tallgrass prairie range in the Osage Cuestas, one of the western portions, of the Central Lowlands physiographic province immediately to the east of the Flint Hills portion of the Central Lowlands. Note: although some of the level III ecosystems for Kansas Chapman et al., 2001). took the names of the physiographic provinces Fenneman, 1938), these two did not coincide completely. For example, Flint Hills Ecoregion, 28a extended eastward into the Osage Cuestas (=Questas) physiographic unit so that the Osage Cuestas Ecoregion, 40b is smaller than the Osage Cuestas physiographic section (comparison of Fenneman, 1938, ps. 614-616 to Chapman et al., 2001). Same situation obtained for level III ecoregions of Oklahoma (Woods et al., 2005) as compared to physiographic units of Fenneman (1938). Source of confusion.

26. Upland tallgrass (bluestem) prairie- Big bluestem was the dominant and prairie dropseed the associate species on this Osage Questas tallgrass prairie range stocked with steers under Intensive Early Stocking. Indiangrass and upland switchgrass were the next two important species (based on both apparent dominance and biomass or herbage production). There were some invaders like Baldwin ironweed, Johnsongrass, curly dock (Rumex crispus), tall fescue, and smooth bromegrass. Silver bluestem and buffalograss were the major native grasses that were in the increaser and invader categories, respectively. Decreaser forbs included Maxmillian sunflower, compass plant, Illinois bundleflower, and leadplant.

Greenwood County, Kansas. Estival aspect, June. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 601 (Bluestem Prairie), generally, or, probably more specifically, SRM 710 (Bluestem Prairie). "Tall-Grass" Series, 142.11, of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Flint Hills Ecoregion, 28a (Chapman et al., 2001).

27. Bluestem prairie grazed by stockers- Tallgrass prairie dominated by big bluestem with prairie dropseed as the associate being grazed by a mixed (both steers and heifers) herd of stocker cattle under Intensive Early Stocking. Note the degree of use on this bluestem-dropseed tallgrass range at beginning of summer (mid-June). Indiangrass and upland switchgrass were the other two of the Four Horsemen of the Prairies grasses that were abundant on this outstanding natural pasture. Little bluestem frequently ranks a distant fourth among the Four Horsemen on tallgrass prairie pastures in the Osage-Flint Hills sections of the Central Lowland grasslands. Illinois bundleflower and leadplant were present but heavily grazed.

The main mid-grass species was sideoats grama; the most common shortgrass species were hairy grama and buffalograss. Mid- and shortgrass species were restricted to localized microsites (perhaps formed partly by spotgrazing). In species composition (density, cover, biomass, etc.) these latter groups constituted nothing even approaching their proportions typical of mixed prairie. A mere listing of species on this form of tallgrass prairie likely would not be different from a species list taken on mixed prairie, except for an occasional species. The difference is in relative proportion of the same, rather than in different, species of grasses. (The swards presented in these slides of tallgrass prairie should be compared to those of mixed prairie in that portion of this publication.)

There were traces of three introduced (agronomic) grasses that would automatically be classified as invaders and that purist prairiemen regard as weeds: smooth bromegrass, tall fescue, and Johnsongrass.

This range had been fired approximately two months prior to this photograph. Tallgrass prairies used for stocker ranges in this "neck of the woods" are traditionally burnt just before green-up each spring, or at least every few years. Otherwise, in instances where cattlemen lease range from local landowners, stocker operators demand a lower pasture rental fee.

Osage Questa sub-unit of the Osage Section of the Central Lowland physiographic province. Limestone is the underlying parent material. Greenwood County, Kansas. Estival aspect, June. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 601 (Bluestem Prairie), generally, or, more specifically, SRM 710 (Bluestem Prairie). "Tall-Grass" Series, 142.11, of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Flint Hills Ecoregion, 28a (Chapman et al., 2001).

28. Steers and heifers grazing tallgrass prairie range in the Osage Questa portion of the once vast bluetem prairie region. Note the thrifty condition and degree of finish on these cattle. They are literally grass-fat which attest to the "strength" (high nutritive value) of this native forage. Degree of use and herbage cover on the land is typical of Intensive Early Stocking which was being used on this range. Close-up of the same range and herd presented in the preceding slide.

Greenwood County, Kansas. Estival aspect, June. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 601 (Bluestem Prairie), generally, or, more specifically, SRM 710 (Bluestem Prairie). "Tall-Grass" Series, 142.11, of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Flint Hills Ecoregion, 28a (Chapman et al., 2001).

29. Ungrazed sward- Upland tallgrass prairie range that had not been grazed by livestock (and barely touched by wildlife) in the current plant-growing season. Furthermore this pasture had not been burnt this spring as shown by numerous dead last year's shoots. This wider view of ungrazed range vegetation presented the physiogonomy, botanical composition, and structure of the vernal society of a Four-Horsemen- of-the-Prairies Flint Hills grassland. This specific grassland community was dominated by an ecotype of upland switchgrass that had unusually wide leaves that formed a "nearly pure" consociation of this species. Big bluestem was the associate species. Redtop panicgrass (Panicum rigidulum) was also present.

Some plants of buckbrush or coralberry (Symphoricarpos orbiculatus).

Greenwood County, Kansas. Mid-June; late vernal aspect. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 601 (Bluestem Prairie), generally, or, more specifically, SRM 710 (Bluestem Prairie). "Tall-Grass" Series, 142.11, of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Flint Hills Ecoregion, 28a (Chapman et al., 2001).

30. Grazed by stocker cattle- A fenceline contrast of Flint Hills tallgrass prairie showing ungrazed sward of highway right-of-way (left) and inside a range grazed by stockers (mostly steers) but not burned this spring (right). Spot grazing and appearance of wold plants was already taking place by late spring and there was still quite a bit of the stocker grazing season remaining (depending, of course, on summer rainfall and consequent soil moisture).

The dominant range plant was switchgrass (an ecotype of the upland form). Big bluestem was the associate species.

31. Grassland diversity- Upland tallgrass prairie of Four Horsemen grasses with switchgrass the most abundant and little bluestem the having least cover and lowest density of the four. This range had not been grazed during the current growing season so that species composition, physiogonomy, and structure of the late vernal society was most visible. Botanical composition was visible in both the general or physiogonomic perspective (first slide) and a close-in "photoplot" view (second slide).

In addition to the tallgrass species there were a number of forbs, including palatable legumes, present. Forb species included wild licorice (Glycyrrhiza lepidota), blue wild indigo (Baptisia australis), leadplant (Amorpha canescens), wild alfalfa or scurfpea (Psoralea tenuifolia) among the legumes and two conspicuous composites, ashy sunflower (Helianthus mollis) and giant or great ironweed (Vernonia crinita).

Chase County, Kansas. Mid-June; late vernal aspect. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 601 (Bluestem Prairie), generally, or, more specifically, SRM 710 (Bluestem Prairie). "Tall-Grass" Series, 142.11, of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Flint Hills Ecoregion, 28a (Chapman et al., 2001).

32. Stockers on tallgrass hills- Okie steers (from about seven hundred to almost eight hundred pounds) on Flint Hills tallgrass prairie range. This was grassland was comprised primarily of the Four Horsemen of the Prairies grasses. Switchgrass was the most abundant plant species while big bluestem was the associate species. The ungrazed range plant community was presented in several slides-photo captions immediately below.

Two behavioral patterns of cattle were evident in these photographs. Cattle, especially younger cattle, are quite curious and will often come right up to visiting humans if they move slowly enough. This is particularily the case if people remain outside the pasture fences, at least until cattle have adjusted to human presence. These three slides were taken with a wide angle (28mm) lens so steers were quite a bit closer than they appeared in photographs. The second behavior shown in these slides was the tendency of cattle newly introduced into a pasture to walk fence lines, the grazable boundaries of the range. Repeated walking of fence boundaries was evident in all three of these slide (and the first of the two-slide set immediately below) by the deep cattle paths cut through the dense, tough prairie sod down to the soil surface.

Tallgrass Prairie National Preserve, Chase County, Kansas. Mid-June; late vernal aspect. K-66 (Bluestem Prairie). SRM 601 (Bluestem Prairie), generally, or, more specifically, SRM 710 (Bluestem Prairie). "Tall-Grass" Series, 142.11, of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Flint Hills Ecoregion, 28a (Chapman et al., 2001).

33. What neither stockers nor fire consumed- Perimeter five-wire fence separating stocker range of Four Horsemen tallgrass prairie which was shown in the preceding three-slide set from highway right-of-way made an exclosure of prairie vegetation on the outside fencerow. This ungrazed (by stocker cattle) range plant community was on the right in the first of these two slides and in fore- and midground of second slide. This particular range (and exclosure) had not been burnt this year (not fired in spring or late winter prior to current warm-growing season) so undecomposed biomass of the previous year was present along with current year herbage.

Switchgrass and big bluestem were dominant and associate plant species, respectively, in the exclosure although range vegetation was nearly a "pure stand" consociation of switcvhgrass. The major forb was Illinois bundleflower (Desmanthus illionensis). Purple prairie-clover (Petalostemon purpurea) was the second-most abundant forb (ie. major forbs were native legumes).

34. In an exclosure- A closer-in view of range vegetation in highway right-of-way (a de facto exclosure) outside the steer range of Four Horsemen tallgrass prairie presented in the three-slide set above the preceding two-slide set (two slide-caption sets above).

This local range plant community was a consociation of switchgrass (upland ecotypes) with Illinois bundleflower, a legume of Caespinoideae subfamily, being the main forb. There were some plants of purple prairie-clover, a papilionaceous legume (Papilionoideae subfamily), was the second major forb. Big bluestem was the associate range range species, but this tallgrass is a "short-shoot" or "culmless" grass that does not elongate its culm until much later in the warm-growing season so that it was not prominent in these two "photoplots". By contrast, switchgrass is a "long-shoot" or "culmed" species (a grass that elongates its culm early in the annual growth cycle) such tht it showed up more conspicuously in these "photoquadrants". Indiangrass and little bluestem, the other two Four Horsemen grasses, are "long-shoot" and "short-shoot" grasses, respectively.

35. Winter sward- Local assemblage of major grass species on an upland tallgrass prairie in the Ozark Plateau. Local dominant species was tall dropseed (Sporobolus asper), seen as the taller, buckskin-colored shoots, with big bluestem (reddish brown patch in right-center of both slides), and Indiangrass (in front and immediately behind the big bluestem). There were also some shoots of broomsedge bluestem (Andropogon virginicus) in background and lower right corner.

This local patch of relict vegetation was in an outside fencerow where lack of grazing defoliation had permitted maximum expression of physiogonomy, structure, and composition of the climax (potential natural) grassland vegetation.

Ottawa County, Oklahoma. Early January; hibernal aspect (dead herbage; dormant phenological stage).

36. Stockers on Osage spring range- Okie steers from Florida on a loamy prairie range site of the bluestem-Indiagrass prairies in the Osage Questas section of the Central Lowlands. Note physiography of the land and height of the tallgrasses after only one month of growth and relatively heavy stocking.

Tallgrass Prairie Preserve (former Chapman-Barnard Ranch), Osage County, Oklahoma, Early May, vernal aspect. FRES No. 39 (Prairie Ecosystem), K-66 (Bluestem Prairie). SRM 710 (Bluestem Prairie). "Tall-Grass" Series, 142.11, of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Flint Hills Ecoregion, 28a (Woods et al., 2005).

37. Greening up and shedding off- Spring in the Osage brings green-up of the perennial grasses (mostly of Andropogoneae and Paniceae tribes) from their root crowns or proaxes (proaxis is the lowest portion of a grass shoot where the nodes/internodes are stacked close atop one another and where shoot emergence takes place; the point of union of shoot and root portions). Meanwhile the buffalo are shedding their heavy winter pelage or “coats” (buffalo hides were known to buffalo runners, the proper name for buffalo hunters, as "“robes"and they called the hair “wool”). The two range sites of Loamy Bottomland and Loamy Upland (for the Osage Questas physiographic unit of the Central Lowlands) are visibly divided by the different shades of green in the newly emerged grasses and grasslike plants on this tallgrass prairie. The plants immediately adjacent to the ephemeral stream are spike rush (Eleocharis spp.). Behind this is a zone of switchgrass distinguishable by last year’s dead stalks. Big bluestem dominates the Loamy Upland range site behind. Note the buffalo wallow (and consequent soil erosion) on the ridgeline. Range students should nevr lose sight of the fact that the ecological interaction known as herbivory includes more than actual defoliation. Trambling and even erosion of bare soil are a natural part of herbiivory as are dunging (a form of both defoliation as well as nutrient recycling), propagule dispersal, etc.

Tallgras Prairie Preserve (former Chapman-Barnard Ranch), Osage County, Oklahoma. Early May, vernal aspect, . FRES No. 39 (Prairie Ecosystem), K-66 (Bluestem Prairie). SRM 710 (Bluestem Prairie). "Tall-Grass" Series, 142.11, of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Flint Hills Ecoregion, 28a (Woods et al., 2005).

38. Indian Territory as the redman knew it- Tallgrass prairie that is a consociation of big bluestem in western Osage Plains (Cherokee Prairie) of northeastern Oklahoma (formerly part of Indian Territory prior to statehood). Other tallgrass species included prairie cordgrass (the local associate species) with lesser amounts of switchgrass and Indiangrass. A locally dominant grass around local depressions (potholes) was longspike tridens (Tridens strictus) which is typically characteristic of mid-sere successional states of tallgrass prairie and tallgrass prairie-oak/hickory savanna (Tyrl et al., 2008, p. 181). Forbs included prairie parsley (Polytaenia nuttallii), the conspicuous umbelliferous plant in foreground of the first slide, and southern wild hyacinth.or Atlantic (prairie) camas (Camassia scilloides).

This meadow had been burnt approximately three months prior to time of photographs. Such prescribed fires encourage big bluestem including sexual reproduction (grain production) as well as growth and vegetative regeneration.

Lone Oak Meadow, Kelly Ranch, Craig County Oklahoma. Mid-May, vernal aspect. FRES No. 39 (Prairie Ecosystem), K-66 (Bluestem Prairie). SRM 710 (Bluestem Prairie). "Tall-Grass" Series, 142.11, of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Central Irregular Plains- Osage Cuestas Ecoregion 40b (Woods et al, 2005).

39. Tallgrass heaven in the spring- A consociation of big bluestem comprised the range plant community of this tallgrass prairie in the Cherokee Prairie of northeastern Oklahoma late spring. Prairie cordgrass was a local associate to even co-dominant in lower-lying microsites of this prairie hay meadow. Longspike tridens was a local dominant around small areas of microrelief that ponded water in wet springs and right after heavy rains (shown below). Switchgrass and Indiangrass were associate species with their relative abundance varying at local scale.

The major forb in this vernal aspect/society was southern wild hyacinth or prairie camas which, with its bright pink petals, set off the prairie vegetation at this season.

The second of these two slides (vertical image) served as a "photoquadrant" of the sward of the vernal society of this tallgrass prairie with an emphasis on prairie camas or wild hyacinth which was a peak, full-bloom stage.

Truly heaven on Earth for a prairieman.

40. Looking into the graass and the sun-View of shoots of big bluestem-dominated sward of tallgrass prairie in western Osage Plains (Cherokee Prairie) in northeastern Oklahoma from a view looking into the bright, spring sun. The immediately preceding four photographs (two sets of two slide-photocaption) and the succeeding slide were of this same tallgrass prairie vegetation (same hay meadow) with sun behind the photographer. By contast, this photograph was takn facing directly into the sun in a cloudless sky.

Direction of light flow can have a major affect of photographs-- for the better or worse depending on what the photographer desired to show or the affect he desired to impart to an image. Photography, like management of ranges, pastures, and forests, will alswys be an art form as well as a science. This into-the-sun shot imparted the brightness of a cloudless spring afternoon on the prairie as well as showing details of foliage that could not be displayed with photographs taken away from direction of direct sunlight.

The tallgrass prairie, and the view of its sward shown here, was a consociation of big bluestem with prairie cordgrass and longspike tridens as local associates to co-dominants around microhabitats of ponded water (see next slide). Trees in distant background were growing along a small prairie stream.

41. Local pothole made a difference- Microrelief on a big bluestem-dominated tallgrass prairie that resulted in a local depression or wet microhabitat that held water during much of the wet months of April, May, and first half of June as well as after heavy rains at other times of the year. This pothole was a unique microenvironment on a predominately upland prairie.

The pothole served as an ideal environment for longspike tridens (the light brown or buff-colored dead grass shoots) and prairie cordgrass, the two co-dominant species of this atypical, local habitat. Although longspike tridens is usually regarded as a species of mid-sere (= an increaser), it is commonly a local climax dominant or co-dominant with other moisture-loving prairie grasses such as prairie cordgrass, eastern gamagrass, and bottomland ecotypes of switchgrass.

42. Grand sweep of the tallgrass prairie- Three landscape-scale views of Osage Cuestas bluestem prairie (the Four Horsemen of the Prairies: big bluestem, little bluestem, , Indiangrass, and switchgrass) with big bluestem the major dominant. Excellent range condition class with both cool-season decreaser grasses as, for example, a lot of Canada or nodding wildrye (Elymus canadensis) and warm-season indicator grasses in addition to the Four Horsemen including lots of prairie dropseed (Sporobolus heterolepis). There was also a wide diversity of prairie forbs including Baldwin ironweed (Vernonia baldwinii), Indian plantain (Cacalia tuberosa), wild alfalfa (Psoralea tenuifolia), and antelopehorn milkweed (Asclepias asperula var. decumbens), and common evening primrose (Oenothera strigosia ssp. canovirens ). There were also several species of umbrella sedge (Cyperus spp.).

These photographs presented the physiography of the southern Osage Cuestas (of the Osage Plains Section in the Central Lowlands physiogrpahic provinc)e and the physiogonomy of tallgrass prairie in early estival aspect as had been modified by moderate stocking of steers which had just been shipped.

Chautauqua County, Kansas. Late June. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 601 (Bluestem Prairie), generally, or, probably more specifically, SRM 710 (Bluestem Prairie). Bluestem "Tall-Grass" Series, 142.11, of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Flint Hills Ecoregion, 28a (Chapman et al., 2001).

43. A treasured meadow- Textbook example of a cherished hay meadow dominated by big bluestem with upland switchgrass as associate species in the Osage Cuestas form of tallgrass parairie. These two photographs were taken at almost the identical global position and within moments of each other in mid-afternoon.. The first of these two photographs was taken under an overcast sky whereas the second photograph was taken just a few minutes later under a full-sun sky. Kodachrome film captured coloration as seen by the human eye so that these two slides showed the difference in color of grassland vegetation as affected by filtered and direct sunlight. A standard, minimum light-filtering skylight was used in front of the wide-angle (28mm) lense.

This same difference in cloudy versus cloudless sky was shown in the next photographs whowing sward of this tallgrass meadow.

Woodson County, Kansas. Late June; early estival aspect. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 601 (Bluestem Prairie), generally, or, probably more specifically, SRM 710 (Bluestem Prairie). Bluestem "Tall-Grass" Series, 142.11, of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40).Central Irregular Plains- Osage Cuestas Ecoregion, 40b (Chapman et al., 2001).

44. Early summer sward- Details of sward of tallgrass prairie with big bluestem as the dominant species and upland switchgrass as associate species. The major forb was Indian plantain with purple prairie clover the second most common forb. These two species represented the Compositae and Leguminosae, respectively, the first and second major families (based on numbers of species and relative abundance as judged by cover, biomass, density, etc.) of forbs on North American grasslands.

The first of these two slides was taken under an overcast sky whereas the second slide was taken just moments later when cumulus clouds that had partially blocked (filtered) sunlight had dissipated. This same contrast was shown in the two immediately preceding photographs. The yellowish tinge to green of foliage in full sun is the same as seen by the human eye. Cloud cover partially blocks some of the natural coloration in living plant tissue. this tends to be more in the yellow and red parts of photosynthetically active radiation of the spectrum (light visible to the human eye).

45. Upland tallgrass prairie- Big bluestem was the dominant and eastern gamagrass was the associate species on this form of bluestem praire. Forbs included leadplant, Illinois bundleflower, purple milkweed (Asclepias purpurea), and the naturalized yellow sweet clover (Melilois officinalis). Even on this virgin tallgrass prairie in "mint condition" there were occasional plants of smooth bromegrass (Bromus inermis) and tall fescue (Festuca arundinacea).

Osage Questa (physiographic sub-province of the Osage Section of the Central Lowlands physiographic province) form of tallgrass prairie. Woodson County, Kansas. Estival aspect, June. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 601 (Bluestem Prairie), more generally, or SRM 710 (Bluestem Prairie). Bluestem "Tall-Grass" Series, 142.11 of Plains Grassland, 142.1 biotic community in Cold Temperate Grassland 142 (Brown et al., 1998, p. 40). Central Irregular Plains- Osage Cuestas Ecoregion, 40b (Chapman et al., 2001).

46. King and First Knight of the Round Table- A hay meadow on an upland form of tallgrass prairie in the Osage Cuestas co-dominated by big bluestem and eastern gamagrass. Very few other grass species were present. The other major graminoid on this meadow was whip or tall nut-rush (Scleria triglomerata) which was an associate species in some local habitats (mirosites). In other local environments the associate species was Helianthus maximiliani, Maximilliian sunflower (eg. immediate foreground, second slide). Other important forbs in this climax grassland were common ragweed (Ambrosia artemisilifolia), leadplant (Amorpha canescens), pale Indian plantain (Cacalia tuberosa), and purple prairie-clover.

Tall or whip nut-rush is a species that this author has found growing on degraded prairie hay meadows as well as on climax or pristine condition tallgrass prairies over much of the humid zone throughout the Ozark Mountains westward through the Osage Plains into isolated parts of the Flint Hills.

Over much of the tallgrass prairie region big bluestem is first in abundance, cover, etc. (as well as palatability to grazing animals) among the Four Horsemen grasses (Weaver and Clements, 1938, p. 520; Phillips Petroleum Company, 1963, p. 9; Stubbendieck et al., 1992, p. 33; Haddock, 2005, p. 281). Big bluestem is thus sometimes regarded as "king of the tallgrass prairie". Big bluestem is justifiably the State Grass of Kansas and Missouri. Eastern gamagrass or corngrass is typically not a dominant species (based on abundance, cover, biomass, etc.) of climax tallgrass prairie communities except on relatively restricted bottomland range sites where it is often co-dominant with prairie cordgrass (Spartina pectinata). (An example of an eastern gamagrass-prairie cordgrass bottomland community was treated immediately below.)

47. More of the King and First Knight- An upland, mesic tallgrass prairie co-dominated by big bluestem and eastern gamagrass. These are two progressively closer-in views of the range plant community introduced in the preceding three-slide side of a prairie hay meadow in the western Osage Plains. The first photograph featured a broad vista of part of the tallgrass meadow dominated by big bluestem with local spots dominated by eastern gamagrass. The second photograph featured an in-your-face view of the foliage within a patch of eastern gamagrass.

This hay meadow consisted of such patches alternatively dominated--almost exclusively--by one or the other of these two decreaser grass species (ie. a mosaic of consociations of big bluestem and of eastern gamagrass alternating back-and-forth). The only other major graminoid was the grasslike species, tall or whip nut-rush. Important forbs included Maximilian sunflower, leadplant, purple prairie-clover, common ragweed, pale Indian plantain, and western yarrow (Achillea langulosa), and some species of goldenrod (Solidago sp.) that was unidentifiable in its present vegetative (pre-sexual) state of phenology.

For whatever reason, prairie cordgrass which is commonly a co-dominant or, at least, an associate with eastern gamagrass, was not present on this mesic upland site. The next series of slide-caption sets presented such an eastern gamagrass-prairie cordgrass prairie community.

48. Now for a bottomland tallgrass community- Within few miles of the mesic, upland tallgrass prairie of big bluestem and eastern gamagrass described in the two preceding slide-caption sets there was an even more mesic and bottomland range site co-dominated by eastern gamagrass and prairie cordgrass. These two "photoquadrants" of ungrazed highway right-of-way introduced the range vegetation of this bottomland prairie community where protection from livestock grazing (ie. pasture fence and a highway formed a de facto exclosure) was a control plot of the climax (potential natural) vegetation.

The large cespitose (bunchgrass) plants were eastern gamagrass, a conspicuous one of which was in late bloom stage in the first "photoplot". The lighter or more pale green (as in the strip in center midground of second "photoplot") was prairie cordgrass that was all in pre-bloom stage..

49. Grazed and (more or less) ungrazed- Three views of a bottomland range site of tallgrass prairie co-dominated by eastern gamagrass and prairie cordgrass with ungrazed highway right-of-way in foreground and prairie grazed by cow-calf pairs (livestock pasture) in background. Details of the grazed prairie (the cattle range) were presented and described below while these three "photo-transects" presented detailsed study of the ungrazed prairie vegetation.

The most glaring difference between prairie vegetation in the cattle range and that of the exclosure (the strip of highway-right-of way land between fence and pavement) was presence of key (= indicator) species of forbs. The three most prominent of these were Maximilian sunflower, annual sunflower (Helianthus annus), and Illinois bundleflower. Plants with large deltoid leaves were annual or common sunflower. Annual sunflower is an extremely variable species, especially phenotypically. McGregor et al. (1986, p. 954) described annual sunflower as "... a huge, polymorphic complex encompassing numerous wild and weedy races...".

50. Look what the cattle did- Stark contrast between a pasture grazed by beef cows and calves (right side of fence) and an ungrazed highway right-of-way (left side of fence) on a bottomland range site of tallgrass prairie in the Osage Cuestas. Selective grazing by cows and calves over countless grazing/warm-growing seasons resulted in reduction of eastern gamagrass and consequent, concomittant increase in cover and biomass (relative abundance) of prairie cordgrass with near-elimination of Maximilian sunflower, annual or common sunflower, and Illinois bundleflower.

The first and last of these forbs have long been recognized as extremely palatable species, being indicator and species on some range sites (Phillips Petroleum Company, 1963, ps. 82, 136). By contrast annual sunflower (in its undomesticated forms) is generally a weedy species usually regarded as an ecological invader. Although annual sunflower is "a classic indicator of disturbed soil" is is also "... palatable and readily sought out by livestock..." (Tyrl et al.2008, p. 347). High palatability of annual sunflower was evident from the contrast presented here in which the grazed pasture served as typical (grazing) management or, in this case, as the control plot while the right-of-way served as an exclosure.

51. Cattle on bottomland tallgrass prairie- General views of a cattle range on a bottomland range site of tallgrass prairie co-dominated by eastern gamagrass and prairie cordgrass The former species was considerably reduced while dominance by the latter species was greatly expanded via selective grazing by cattle over a number of years. It was shown and explained in the preceding slide and caption that cover and general abundance of Maximilian and annual ssunflowers along with that of Illinois bundleflower was also reduced by cattle grazing.

These changes is species composition--including the palatable though generally weedy annual sunflower--constituted overgrazing which by definition is grazing-induced departure from the climax plant community to a seral stage (or, actually, movement of the range vegetation from any higher successional stage to a lower successional state) so as to result in range deterioration or degradation. This change in species composition is retrogression, generally the opposite direction of plant community dynamics from that of plant succession. Overgrazing (an overgrazed vegetational state) is the result of long-tern overuse, forage harvest in excess of sustained yield (= overstocking; harvesting range feed resources beyond grazing capacity of the range).

Even though there appeared to still be "planty of feed" with a lot of "high grass" (and made up of tallgrass species) this range was overgrazed. Admittedly departure from climax vegetation was not severe and there was still substantial cover of eastern gamagrass and no apparent soil erosion. Nonetheless grazing management .had not been proper. There was clear departure from the climax vegetation which, in the case of this range site, was the most productive for cattle (feeder calf) production. Eastern gamagrass is much superior to prairie cordgrass in palatability, biomass production, and nutritive value of forage. That is why beef cattle had selectively eaten gamagrass in preference to cordgrass. This pattern was even more pronounced in regards to near-elimination of palatable composite and legume forbs.

Associated with slight overgrazing was even more fragrant underburning, inadequate use of prescribed fire. Together these two malpractices (at very least, less-than-ideal management practices) had permitted woody invasion of this grassland by green ash (Fraxinus pennsylvanica), a native tree species kept to minimal cover on virgin tallgrass prairie by frequent range fires. Green ash varied in size from large seedlings to mid-sized saplings in fore- and midground to larger adult trees in background of these photographs. (Size of large seedlings was more apparent relative to the Angus cattle in the next set of three slides.)

52. Some of the ice cream species, along with an invading one- A cattle pasture on a bottomland range site of tallgrass prairie co-dominated by prairie cordgrass and eastern gamagrass showing reduction of gamagrass, concomittant increase of cordgrass, and invasion by green ash due to some overgrazing and underburning of this range. In each of these three photographs there were small green ash as tall as the back or shoulders of adult Angus cows. Green ash is a native tree of this range plant community, but in virgin (climax) state green ash is a minor component of the range vegetation, ash being kept as an incidental species by recurrent prairie fires and grazing that permits the two climax decreaser grasses to "keep out" (or, at least, to low cover) what constitutes a woody invader of this grassland.

The case could be made that with considerable cover of eastern gamagrass--though more as an associate than co-dominant species--remaining the degree of overgrazing was not critical or, perhaps more accurately, had not yet reached the point that impacted management of this range in a practical way. That argument might well be valid with regard to eastern gamagrass, but such argument fails--fails miserably--with regard to dramatic reduction (indeed almost complete elimination) of climax (decreaser) forbs such as Maximillian sunflower and Illinois bundleflower. Instead a major forb on the cattle range was pale Indian plantain, a good specimen of which was in lower right corner of second slide.

Height of eastern gamagrass could be guaged by noting that some of its shoots were as tall or taller than calves and almost as tall as invading young green ash..

Eastern gamagrass on some range sites is an ice cream species, an unusually palatable species consumed first by range animals so that plants of such species are commonly overused and, thus, difficult to maintain even under proper grazing management (Kothmann, 1974). On this tallgrass prairie bottomland range site where eastern gamagrass is the climax co-dominant with prairie cordgrass, eastern gamagrass is not an ice cream species. Rather, on this bottomland range site eastern gamagrass is a key species which is by definition 1) a species which is of such importance that it has to be managed for (ie, maintained on the range) and 2) a feed species the use of which serves as an indicator of degree of use of other range species (Kothmann, 1974).

Successional and management status of any range plant species is range site-specific.

53. Tallgrass stocker range- Another range of the Four Horsemen of the Prairies--big bluestem, Indiangrass, switchgrass, little bluestem--being grazed by about five-weight (500 pound) steers. These landscape-scale views presented the topography as well as physiogonomy and structure of tallgrass prairie at its late vernal aspect in the Osage Questas of the Central Lowlands physiographic province. Other grasses besides the "big four" dominants included prairie cordgrass (Spartina pectinata), Junegrass, Virginia wildrye, and the native annual Agrostis hyemalis, winter bentgrass. Major forbs included Baldwin ironweed (Vernonia baldwinii), catclaw sensitivebriar, wild alfalfa or slender scurfpea (Psoralea tenuiflora), Maxmillian sunflower, annual fleabane (Erigeron annuus), largeflower tickseed (Coreopsis grandiflora), and pale purple coneflower (Echinacea pallida). Shrubs were essentially non-existent as this range was burned almost every spring.

The soil was Kenoma silt loam, a fine, smectitic (montmorillonitic) thermic Vertic Argiudoll (Soil Conservation Service, 1990). Range site was Clay Upland.

Coffey County, Kansas. Mid-June. (late vernal aspect). FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 601 (Bluestem Prairie), more generally, or SRM 710 (Bluestem Prairie). Bluestem "Tall-Grass" Series, 142.11 of Plains Grassland, 142.1 biotic community in Cold Temperate Grassland 142 (Brown et al., 1998, p. 40). Central Irregular Plains- Osage Cuestas Ecoregion, 40b (Chapman et al., 2001).

54. Grasses, legumes, and composites- Sward of a tallgrass prairie range showing array of species in the three major plant families of this vegetation. This was just outside the fence of the stocker range introduced immediately above. In this photograph of ungrazed range vegetation the following species were found 1) grasses: big bluestem, little bluestem, Indiangrass, tall dropseed, Junegrass, Virginia wildrye, winter bentgrass, plus naturalized smooth brome and tall fescue (domestic pasture grasses); 2) legumes: wild alfalfa or slender scurfpea and catclaw sensitivebriar; and 3) composites: Maxmillian sunflower, annual fleabane, and largeflower tickseed.

These same species were also inside the fenced pasture though wild alfalfa and Maximillian sunflower had been grazed down considerable so as to be less obvious in the range vegetation.

Coffey County, Kansas. Mid-June. (late vernal aspect). FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 601 (Bluestem Prairie), more generally, or SRM 710 (Bluestem Prairie). Bluestem "Tall-Grass" Series, 142.11 of Plains Grassland, 142.1 biotic community in Cold Temperate Grassland 142 (Brown et al., 1998, p. 40). Clay Upland range site. Central Irregular Plains- Osage Cuestas Ecoregion, 40b (Chapman et al., 2001).

55. Grazed and ungrazed- Fenceline contrast between Four Horsemen-dominated tallgrass prairie (big bluestem was the number one dominant) steer range and ungrazed vegetation just across the pasture fence. The exclosure (ungrazed) vegetation was in the foreground (in front of the fence) in the first slide and to the right of the fence in the second slide. This tallgrass prairie was stocked fairly heavily at something like Intensive Early Stocking. This was private range managed commercially so the exact features of Intensive Early Stocking were not followed as would have been the case on an experimental range or even with xtra-conscientious commerical management. Grazing treatment was similar enough for discussion.

Big bluestem was the overwhelming dominant and prairie cordgrass and switchgrass were the local associate species on this mesic lower area of a Loamy Upland range site. At least that was the species composition on the ungrazed side of the fencerow. Unfortunately that was not the same situation immediately inside the steer-grazed range where tall fescue had invaded and was in fierce competition with the native grasses. The naturalized Eurasian tall fescue is a cool-season species whereas the native dominants are warm-season panicoid or eragrostoid species. Steer ranges like this one are grazed only during the warm growing season so that much of the early growth of the endophyte-hosting, grazing-tolerant tall fescue has been made prior to cattle grazing. Tall fescue is much less palatable than the native tallgrasses so that under warm-season grazing tall fescue has a decided advantage over big bluestem, switchgrass, prairie cordgrass, and other indigenous species. This author could not find any tall fescue on the outside of the fencerow. Apparently this agronomic pasture grass cannot compete effectively with the tallgrass species except under grazing where there is clearly selective grazing of the native tallgrasses in preference to the endophyte-infected and generally low-palatability tall fescue. Under spring to early summer grazing the invasive tall fescue benefits from heavy grazing, at least during that time period.

Conversely, if cattle are shipped early enough in the summer--as under proper administration of Intensive Early Stocking--the native range plants would generally have ample time to recover photosynthetic biomass and restore food reserves in root crown and roots to carry out respiration through their dormant-season. However, tall fescue would be ungrazed (other than by wildlife) during its entire autumn and late winter-early spring periods of regrowth and new growth, respectively. If this prairie owner is not careful he will end up owning just another ordinary pasture of tall fescue and loose a treasured (it should be anyway) tallgrass prairie. The enemy is looming on the prairie horizon.

There was an atypically higher density of Baldwin ironweed on this prairie, both inside and outside the fence. There was obviously greater cover and density of ironweed on the steer range, but this conspicuous difference was not a glaring one. This tallgrass prairie range was not in any danger of becoming a weed patch--other than of tall fescue which was the most threatening noxious plant on this rangeland.

In essence, most of the difference--at this point in time--in grazed versus ungrazed vegetation was degree of use not species composition.

Coffey County, Kansas. Mid-June. (late vernal aspect). FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 601 (Bluestem Prairie), more generally, or SRM 710 (Bluestem Prairie). Bluestem "Tall-Grass" Series, 142.11 of Plains Grassland, 142.1 biotic community in Cold Temperate Grassland 142 (Brown et al., 1998, p. 40). Clay Upland range site. Central Irregular Plains- Osage Cuestas Ecoregion, 40b (Chapman et al., 2001).

56. Growth by late spring- Two "photoquadrants" of ungrazed sward of tallgrass prairie (big bluestem clearly dominant) just outside a fairly heavily grazed stocker range. Details of the exclosure vegetation presented in the two preceding slides. Although big bluestem furnished most of the herbage seen in the first slide there was considerable cover of swithcgrass and prairie cordgrass. There was also some wild alfalfa or slender scurfpea in the first "photoplot". Note in the second slide the already dead shoots of the cool-season Junegrass that had just entered dormancy. The larger, prominently wider leaves behind the Junegrass were those of switchgrass whereas the narrower leaves and generally lower shoots were of big bluestem.

Coffey County, Kansas. Mid-June. (late vernal aspect). FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 601 (Bluestem Prairie), more generally, or SRM 710 (Bluestem Prairie). Bluestem "Tall-Grass" Series, 142.11 of Plains Grassland, 142.1 biotic community in Cold Temperate Grassland 142 (Brown et al., 1998, p. 40). Clay Upland range site. Central Irregular Plains- Osage Cuestas Ecoregion, 40b (Chapman et al., 2001).

57. Subtle but real- Fencerow relict vegetation (first photograph) and heavily grazed to overgrazed tallgrass prairie range (second photograph). Range vegetation was straight across a highway from the big bluestem-Indiangrass-switchgrass prairie range described immediately above. The first slide was taken from vantage point of the fencerow looking into a heavily grazed to an overgrazed range stocked with steers during the spring-summer growing period (and over course of many years). Range vegetation on the exclosure (outside the fence on road right-of-way) included eastern gamagrass (the immense clump of giant grass with fully developed inflorescences), big bluestem, switchgrass, and prairie cordgrass. (This range plant community was growing in a transition between lowland and upland range sites.) Also present on the "protected" side of the fencerow was the agronomic (introduced) forage legume, crown vetch (Coronilla varia), which frequently occurs as an adventive or naturalized species. Ohio spiderwort (Tradescantia ohiensis) was also growing to large size on the outside fencerow (right-rear of eastern gamagrass). This member of the dayflower family (Commelinaceae) was not found on the stocker range leading one to suspect that steers had found any spiderworts at least somewhat appentizing or that stocking rate was so high relative to available forage that even unpalatable species were grazed (heavily of course).

Vegetation of the steer pasture was completely devoid of eastern gamagrass (it had been grazed out by stockers over the years) and there was as much (or more) tall dropseed (Sporobolus asper) and poverty dropseed (S. vaginiflorus) than big bluestem. Cover of switchgrass and prairie cordgrass was about the same on both sides of the fence, but Virginia wildrye, the dominant cool-season grass, was either missing or grubbed so close to the soil surface that eyes of an experienced prairieman could not find any. The most abundant forb on this heavily stocked range was Baldwin ironweed as compared to Maximillian sunflower immediately outside the range (see the next two photographs). There were also some plants of annual fleabane on the steer range.

These differences in range vegetation between heavily grazed (by steers) and ungrazed (by livestock and minimally by wildlife) tallgrass prairie were all signs of overgrazing (= long-term overuse resulting in retrogression of the range plant community). Eastern gamagrass, which would have been a "snap" to find--had it existed--on the stocker range given the huge leaves of this species, is difficult to maintain on ranges because it is often an ice cream species. Ice cream species are those that are readily grazed out on ranges due to their extreme palatability and/or limited tolerance to grazing. Loss of ice cream species from the range is often regarded as "collateral damge", unavoidable demise of a species under realistic grazing management (species not worth the cost, effort, concern, etc.). Ice cream species are analogous to sacrifice areas. Specifically, stocking rates and degrees of use would have to be lighter than those necessary for sustained yield on Good to Excellent condition ranges and those that maximize longterm profit from the forage resource. Eastern gamagrass is clearly an ice cream species on certain range sites and when it is a minor member of the range plant community (low proportions of relative cover, biomass, plant density, etc.) or not a key species. That stated, this rangeman has seen numerous tallgrass prairie cattle ranges in the Flint Hills-Osages Cuestas-Chautauqua Hills Region where large plants of eastern gamagrass were present even after years or decades of cattle grazing. On this part of the stocker range where prairie cordgrass and bottomland switchgrass were major species, eastern gamagrass was most probably a realistic persistent species and not an ice cream species. In fact, it is more likely that eastern gamagrass should be a key species to which degree of use and continuing presence in the range plant community would serve as an indication of proper grazing management.

This range was not receiving the respect it should have had. Grassland like this should be cherished as priceless because it is. Incidentially, even the steers were wild in this pasture which is a "dead giveaway" that husbandry is held in low esteem by this steerman and/or landowner. On numerous occasions this photographer has had stocker cattle come right up to the fence to size up him up. They were curious. Wild cattle and degraded ranges are highly correlated. One very good management practice on this range was annual to biennial burning.

Coffey County, Kansas. Mid-June. (late vernal aspect). FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 601 (Bluestem Prairie), more generally, or SRM 710 (Bluestem Prairie). Bluestem "Tall-Grass" Series, 142.11 of Plains Grassland, 142.1 biotic community in Cold Temperate Grassland 142 (Brown et al., 1998, p. 40). Clay Upland range site. Central Irregular Plains- Osage Cuestas Ecoregion, 40b (Chapman et al., 2001).

58. Late spring vestment- Details of the sward of upland tallgrass prairie co-dominated by big bluestem and en upland ecotype of switchgrass. The associate species was the cool-season Virginia wildrye. The major forb was Maximillian sunflower (numerous plants were featured prominently in the second photograph). There was also some tall dropseed, wild alfalfa, and annual fleabane in this range vegetation which was just outside the fence of the heavily grazed stocker range described immediately above. This exclosure (outside fencerow) vegetation was representative of the climax vegetation on upland sites just prior to onset of summer and rapid growth/phenological development. This was the vernal society of tallgrass prairie in Excellent range condition class.

Coffey County, Kansas. Mid-June. (late vernal aspect). FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 601 (Bluestem Prairie), more generally, or SRM 710 (Bluestem Prairie). Bluestem "Tall-Grass" Series, 142.11 of Plains Grassland, 142.1 biotic community in Cold Temperate Grassland 142 (Brown et al., 1998, p. 40). Clay Upland range site. Central Irregular Plains- Osage Cuestas Ecoregion, 40b (Chapman et al., 2001).

59. A little lower down on the range- Slightly lower elevation of tallgrass prairie in a transition between upland and lowland range habitats on the Cherokee Prairie. This was a lower-lying portion of the same stocker cattle range introduced above. Grassland presented here was a three-dominant "botanical mix" of big bluestem, switchgrass (bottomland ecotypes), and prairie cordgrass. It was explained above that eastern gamagrass was a likely member of this range plant community, but no plants of that "grand dad grass" were in this local grassland vegetation that had for years been subjected to summer grazing by stockers.

The first of these two photographs (both taken with a 28mm lens) showed steers starting to mill on being approached (from a distance of roughly 75-80 yards) while the second photograph caught the steers as they broke from the tight herd, high-tailed, and bolted. "Ghost riders in the sky" swung by to try in eternal futility to catch this "Devil's herd" as was foretold by the famed western song writer Stan Jones.

Coffey County, Kansas. Mid-June. (late vernal aspect). FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 601 (Bluestem Prairie), more generally, or SRM 710 (Bluestem Prairie). Bluestem "Tall-Grass" Series, 142.11 of Plains Grassland, 142.1 biotic community in Cold Temperate Grassland 142 (Brown et al., 1998, p. 40). Loamy Lowland range site. Central Irregular Plains- Osage Cuestas Ecoregion, 40b (Chapman et al., 2001).

60. A little lower (and deeper) yet- Cante-cornered and about 150 yards away from the big bluestem-switchgrass-prairie cordgrass prairie range just presented immediately above there was a prairie slough in an adjacent pasture in which the natural vegetation included 1) wet prairie along an ephemeral stream and 2) marsh in the temporarily flooded backwater area (localized floodplain) of the stream. This slide presented the wet prairie along the stream. It was a consociation of prairie cordgrass with rufous bulrush (Scirpus pendulus) as the associate species. The wet prairie was closer to the channel of the ehemeral stream where surface water and soil water drained away faster than in the backwater habitat that was created by a previously eroded, ox bow depression. On that lower, wetter range environment rufous bulrush predominated and prairie cordgrass assumed associate status. Another zone of wet prairie formed on the outer perimeter of the bulrush-dominated marsh so that this tule wetland was surrounded or situated between two strips of wet prairie. On this outer and somewhat less mesic wet prairie indigobush (Amorpha fruticicosa) was the associate to local co-dominant with prairie cordgrass.

The marsh was presented below.

The conspicuous white-flowered forb was the composite, Iindian plantain, which was featured earlier in this chapter.

Coffey County, Kansas. Mid-June. (late vernal aspect). No. 41 (Wet Grasslands Ecosystem). K-65 (Northern Cordgrass Prairie). No SRM. Closest unit in Brown et al. (1998, p. 45) was Cold Temperate Grassland 142., Plains Grassland 142.1 under which there should have been a Cordgrass Series at, say, 142.14.Loamy Lowland range site. Central Irregular Plains- Osage Cuestas Ecoregion, 40b (Chapman et al., 2001).

61. Foundation of wet tallgrass prairie- Profile of the soil in a drainage in tallgrass prairie that formed a slough through which communities of wet praire and marsh had developed in the Cherokee Prairie. This soil was the Verdigris series, fine-silty, mixed, superactive, thermic Cumulic Hapludolls (Soil Conservation Service, 1990). Verdigris soils developed in silty alluvium and are generally very deep yet well drained having formed in landforms of floodplains to local drainage channels such as that shown here.

Range site of a Verdigris is Loamy Lowland. This is one of the most productive range sites in this region of tallgrass prairie. Unfortunately, the color of the soil was not accurately captured by the scanner (Epson Perfection 700) even though Kodachrome 64 film got the color perfectly.

Coffey County, Kansas. Mid-June.

62. Slough in tallgrass prairie- Contact of wet prairie and marsh (first slide) and wetland vegetation of a rufous bulrush-dominated marsh (second slide) that developed in a slough along an ephemeral stream channel (wet prairie vegetation) and on a ox bow depression formed outward from (behind) the channel (bulrush or tule marsh). The first view (slide) was at edge of the marsh with the wet prairie behind while the second slide showed the marsh vegetation of rufous bulrush (dominant) and prairie cordgrass (associate). There was also at least one species each of Carex and Cyperus, neither of which was identifiable at their current stages of development.

Coffey County, Kansas. Mid-June. (late vernal aspect). FRES No. 41 (Wet Grasslands Ecosystem). K-42 (Tule Marshes). No SRM. Closest unit in Brown et al. (1998, p. 45) was Plains Interior Marshland 242.3 under which there should have been a Bulrush or Tule Series at, say, 242.33. Loamy Lowland range site. Central Irregular Plains- Osage Cuestas Ecoregion, 40b (Chapman et al., 2001).

63. Thrive on fire and semi-wet feet- Range vegetation of wet prairie on which prairie cordgrass was dominant and indigobush was associate to local co-dominant. This was an outer zone of wet prairie that developed along the margin of a marsh dominated by rufous bulrush with prairie cordgrass as associate species on that wetland. The soil of this wet prairie was slightly better drained than that of the marsh which enabled the shift between rufous bulrush and prairie cordgrass and permitted survival of the shrub, indiobush. Bulrush did best when its roots stayed in mud where it could out-compete prairie cordgrass whereas cordgrass gained the competitive edge on less wet ground where it gained a "woody pardner".

All three of these prairie plants do quite well under periodic burning. This range had been fired about two and a half months prior to time of these photographs so that all shoot were of present season's growth. Frequent burning might harm indigobush under certain conditions, but this particular range was burnt about every year (sometimes every other year) and this woody legume was still putting forth vigorous resprouts.

Coffey County, Kansas. Mid-June. (late vernal aspect). FRES No. 41 (Wet Grasslands Ecosystem). K-65 (Northern Cordgrass Prairie). No SRM. Closest unit in Brown et al. (1998, p. 45) was Cold Temperate Grassland 142., Plains Grassland 142.1 under which there should have been a Cordgrass Series at, say, 142.14. Loamy Lowland range site. Central Irregular Plains- Osage Cuestas Ecoregion, 40b (Chapman et al., 2001).

Great Plains form of tallgrass prairie- Except for the sand dunes and sandhills forms of postclimax tallgrass the ultimate expression of North American tallgrass prairie reaches its western limits at the eastern perimeter of the Great Plains physiographic province. Part of this is the Plains Border, including the Smoky Hills (Fenneman, 1931, ps. 25-27).

64. About as far west as it goes- Another treasured prairie hay meadow with big bluestem, upland switchgrass, and tall dropseed (Sporobolus asper) with nodding or Canada wildrye as the major cool-season grass species. This jewel supported such prairie legumes as purple prairie clover (Petalostmon purpureum), Maximillian sunflower (Helianthus maximiliani), blackeyed Susan (Rudbeckia hirta), leadplant, wild alfalfa, and Illinois bundleflower.

As if the "mint condition" condition of this lovely meadow was not enough the clinching feature about this native range vegetation was that it was in the some of the more western margins of tallgrass paririe specifically the Plains Border section of the Great Plains physiographic province (Fenneman, 1931, ps.25-27).

Harvey County, Kansas. Late June; early estival aspect. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 601 (Bluestem Prairie), generally, or, probably more specifically, SRM 710 (Bluestem Prairie). Bluestem "Tall-Grass" Series, 142.11, of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40).Central Great Plains- Wellington-McPerson Lowland Ecoregion, 27d (Chapman et al., 2001).

65. The Missouri Natural Areas Committee recognized 14 "prairie natural communities". These were based on a hypothetical topographic-geologic transect, but this relief alignment was itself based on a hypothetical chresard profile (Nelson, 1987, ps. 77-80). This was a generic, theoretical soli catena (catena was defined and illustrated in a succeeding photograph and caption). Soil was not the basis of the types or kinds of prairie other than delineation of alluvium and loess and parent material was not a factor except to show depth to limestone or chert, sandstone, or sand. As such these were not the equivalent of range sites. The Soil Conservation Service did not designate range sites in Missouri. In fact, the SCS in Missouri did not even speak to the matter of range or rangeland The next three slides show examples of some kinds of prairies in Missouri and an adjacent county in Kansas based on the Missouri Natural Areas Committee System.

The 14 Missouri prairie communities were: 1) Dry prairie, 2) Dry-mesic prairie, 3) Mesic prairie, 4) Wet-mesic prairie, 5) Wet prairie, 6) Dry limestone/dolomite prairie, 7) Dry-mesic limestone/dolomite prairie, 8) Dry chert prairie, 9) Dry-mesic chert prairie, 10) Dry sandstone/shale prairie, 11) Dry-mesic sandstone/shale prairie, 12) Dry sand prairie, 13) Dry-mesic sand prairie, and 14) Hardpan prairie.

66. Hardpan Prairie- Little bluestem and upland switchgrass dominate this shallow upland range site in the bluestem prairie region. Prairie dropseed is the main associate but Indiangrass and big bluestem are prominent and contribute substantially to cover and biomass. There is a "healthy"array of forbs, mainly composites. The four visibly prominent purple inflorescences are of showy or elegant gayfeather (Liatris elegans). This delightful prairie has been used as a prairie hay meadow for years. Although tallgrass prairie and oak-hickory prairie savanna are the climatic climax for this region the unique species composition of this grassland is edaphic. A claypan overlaying a seam of coal causes a perched water table of acidic groung water and a soil pH of 4.7. This combination favors prairie dropseed, especially, and little bluestem, secondly, rather than the regional dominants of big bluestem and Indiangrass. The shallow but mesic soil and perched watertable explains upland switchgrass. This form of tallgrass prairie is in the Ozark Plateau or Ozark Highlands physiographic province.

Stoney Point Prairie, Dade County, Missouri. Estival aspect, July. FRES No. 39 (Prairie Ecosystem), K-66 (Bluestem Prairie), Hardpan Prairie, SRM 710 (Bluestem Prairie) and SRM 802 (Missouri Prairie). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Chapman et al., 2002).

67. The "Sea of Grass" is far from monotonous- A landscape-scale scene of tallgrass prairie the Cherokee Prairie Region (the Cherokee Lowlands section of the Central Lowlands physiographic province) may look like sameness or monotony to the "unth degree" to newcomers, but to trained eyes of prairiemen it is anything uniform.

The vast interior of continents was the birthplace of the zonal concept. In this context zonal refers or is applied to "features (eg. soils and vegetation) characteristic of a particular region that is approximately bounded by lines of latitude (ie. a region lying parallel to the equator" (Allaby, 1998) This is the usage that was the basis of newer conceptual views of large spatial scale ecology such as ecoregion (= ecosystem region) as applied in Ecosystem Geography (Bailey, 1995, 1996, 1998, 2002). The earliest application of the zonal perspective to soils has been traced to formal usage in soil classification systems by Russian soil scientists especially Dokuchaev (see for eg. Baldwin et al. in United States Department of Agriculture, 1938, p. 980; Fanning and Fanning, 1989, ps. 141-149 passim). Zonal was basis of the first effort at a comprehensive soil system in the United States, the organization of which was into zonal, intrazonal, and azonal soils (Soil Classification in United States Department of Agriculture, 1938, ps. 979-1001).

In the Bailey (1995, 1996, 1998) Ecosystem Geography view the intermediate spatial heirarchial unit is the landscape mosaic, or simply landscape, "a geographical group of site-level ecosystems" in which site is the range site level (Bailey, 1996, ps. 22-25, 169). Some authors recognized the catena unit as another spatial unit in the heirarchy intermediate between landscape and range site. Archer and Smeins in Heitschmidt and Stuth (1991, ps. 110-112) applied catena in this way: "A catena is comprised of linked ecosystems. The landscape is a hierarchial level comprised of catenas".

Catena in this scheme is obviously the catena used in Soil Science. The Soil Science Society of American (2001) defined catena distinguised it from toposequence.Catena was defined as "a sequence of soils of about the same age, derived from similar parent material, and occurring under similar commatic conditions, but having different characteristics due to variation in relief and in drainage" while a toposequence was "a sequence of related soils that differ, one from the other, primarily because of topography as a soil-formation factor".

Across the portion of landscape mosaic of tallgrass prairie presented above there were probably two or three catenas. There were innumerable hypothetical toposequencial transects that could be measured across the grassland ecosystem (or, depending on spatial scale of study, ecosystems), but this was a catena not a toposequence because drainage regimes were considerably different on the various range sites comprising this landscape.

Using the "prairie natural communities" catena sequence of the Missouri Natural Areas Committee (Nelson, 1987, ps. 77-80) that were listed above there were three Missouri prairie communities across the tallgrass prairie landscape shown in this photograph: 1) Mesic, 2) Wet-Mesic, and 3) Wet Prairies.

Major grasses were the Four Horseman of the Prairies species, prairie dropseed, and prairie cordgrass (Spartina pectinata). Composites were by far the major forbs. The most conspicuous at time of photograph was prairie gayfeather or prairie blazing star (Liatris pycnostachya), a specimen of which was featured in left foreground.

This general geologic region is in the Osage (= Osage Plains) section of the Central Lowland physiographic province (Fenneman, 1938, ps. 455, 605-630 passim), most specifically the Cherokee (or Nevada) Lowlands of the Osage section (Fenneman, 1938, p. 612-613). This region was unglaciated. It is bounded to the southeast by the Ozark Plateaus (= Ozark Uplands) physiographic province and to the west by the Osage Questas section of Central Lowlands. Osage Plains Natural Division (Nelson, 1987, ps. 3 [2-5], 77).

Prairie State Park, Barton County, Missouri. July. Estival aspect. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie), Mesic Prairie, Wet-Mesic Prairie, and Wet Prairie communities. SRM 710 (Bluestem Prairie) and SRM 802 (Missouri Prairie). Central Irregular Plains- Cherokee Plains Ecoregion, 40d (Chapman et al., 2002).

68. Patchwork of tallgrass prairie communities- A vegetational mosaic had developed on this slope of tallgrass prairie in the Cherokee Lowlands physiographic province. Vegetation in foreground (a draw or depression at foot of hill slope) comprised a Wet Prairie community dominated by Carex species with prairie cordgrass and bottomland switchgrass being associates. Background vegetation consisted of two subforms of Wet-Mesic Prairie: a drier subform dominated in roughly equal portions by big bluestem, Indiangrass, and paririe dropseed (left background) and a more mesic subform that was a consociation of prairie cordgrass (right background).

This pattern of different tallgrass prairie plant communities at small range site-scale was result of drainage that followed topographic patterns. For those who can truly see (ie. comprehend or understand) and describe vegetation, tallgrass prairie is often extremely varied over relatively small spatial scale. Tallgrass prairie vegetation is far from homogenous.

Prairie State Park, Barton County, Missouri. July. Estival aspect. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). Wet Prairie and Wet-Mesic Prairie communities. SRM 710 (Bluestem Prairie) and SRM 802 (Missouri Prairie). Bluestem "Tall-Grass Series, 142.11 of Plains Grassland biotic community, 142.1, of Brown et al. (1998, p. 40) as was explained and repeated here for reader convenience. Central Irregular Plains- Cherokee Plains Ecoregion, 40d (Chapman et al., 2002).

69. Consociation amid complexity- A natural single species-stand of prairie cordgrass had developed on a mid-slope above a wet (frequently ponded) depression (see immediately preceding slide). This stand of cordgrass might have been a colony of one genotypic plant rather than a population of several genetic individuals of prairie cordgrass. It was as uniform a crop as any farmer could dream of, but this small "patch" of cordgrass was within (inside of) a larger and diverse range plant community of tallgrass grassland.

Prairie State Park, Barton County, Missouri. July. Estival aspect. FRES No. 39 (Prairie Ecosystem). K-66 (Bluestem Prairie). Wet-Mesic Prairie community. SRM 710 (Bluestem Prairie) and SRM 802 (Missouri Prairie). Central Irregular Plains- Cherokee Plains Ecoregion, 40d (Chapman et al., 2002).

70. More complexity and species richness in the resources-rich habitat of a tallgrass prairie range- Wet Prairie community (foreground) consisting of several sedge (Carex) and umberella or flat sedge (Cyperus) species was in the foreground of this photograph. The two conspicuous grasslike plant species in the foreground were the umberella sedge, also known as green or marsh flatsedge(Cyperus virens= C. pseudovegetus), smaller plants, and straw-colored flatsedge (C. strigosus), larger and taller plants. The prominent white-flowered forb was common mountain mint (Pycnanthemum virginianum).

Mesic-Wet Prairie community was in background. Dominated by prairie cordgrass with big bluestem as associate. The Wet Prairie and Mesic-Wet Prairie communities were distinct and without apparent transition zones between them. Likewise there was no prominent transition between Mesic-Wet Prairie communities and Mesic Prairie communities upslope from them except for increasing predominance of big bluestem and sudden appearance of high cover of Indiangrass.

Prairie State Park, Barton County, Missouri. July. Estival aspect. FRES No.. 39 (Prairie Ecosystem). K-66 (Bluestem Prairie). SRM 710 (Bluestem Prairie). Wet prairie and Wet-Mesic Prairie communities. SRM 710 (Bluestem Prairie) and SRM 802 (Missouri prairie). Central Irregular Plains- Cherokee Plains Ecoregion, 40d (Chapman et al., 2002).

71. Mesic and Wet-Mesic Prairie- This tallgrass prairie in the regional Cherokee Prairie is in the Cherokee Lowlands subunit of the Central Lowlands. This physiographic unit joins the Ozark Plateau immediately to its east. Two of the Missouri kinds or categories of prairie (“prairie natural communities”) are seen here. Overall, big bluestem is the dominant species but in wet depressions of the Wet-Mesic Prairie form species of Carex and Scirpus dominate. On raised, drier microsites of the Mesic Prairie needle-and-thread is common (visible as a purplish color). At this early summer season the short-shoot big bluestem has not elongated its culm so the tallgrass physiogonomy is not apparent and the grassland instead resembes a mid-grass or mixed praririe. Switchgrass is present only as an associate on this mesic to hydric water regime because by mid-summer to autumn the soil is typically relatively dry reflecting the precipitation pattern (ie. June is the wettest month of the year and the soil is driest by August or September).

Crawford County, Kansas. Late vernal to early estival aspect, June. FRES No. 39 (Prairie Ecosystem), K-66 (Bluestem Prairie), Mesic and Wet-Mesic Prairie, SRM 710 (Bluestem Prairie) and SRM 802 (Missouri Prairie). Central Irregular Plains- Cherokee Plains Ecoregion, 40d (Chapman et al., 2002).

72. Dry-Mesic Chert and Dry Chert Prairie- This tallgrass prairie is in close proximity to the Mesic and Wet-mesic kinds of prairie in the previous slide and is also part of the former magnificant regional Cherokee Prairie which joins (what is left of it) the magnificant Flint Hills physiographic province and its famed bluestem range. This is also a Four Horsemen of the Prairies tallgrass form and big bluestem is also dominant overall, but prairie dropseed, little bluestem, and needle-and-thread can individually or collectively dominate locally. The foreground is the dry-mesic chert form while the background is the dry chert form.

Prairie State Park, Barton County, Missouri. Late vernal to early estival aspect, June. FRES No. 39 (Prairie Ecosystem), K-66 (Bluestem Prairie), Dry-Mesic Chert and Dry Chert Prairie. SRM 710 (Bluestem Prairie) and SRM 802 (Missouri Prairie). Central Irregular Plains- Cherokee Plains Ecoregion, 40d (Chapman et al., 2002).

73. Species-rich tallgrass prairie range community- This Mesic Prairie community in the Cherokee Prairie Region was dominated by the Four Horseman of the Prairies species (big bluestem and Indiangrass were prominent) plus prairie dropseed. Eastern gamagrass was also locally abundant. Most notable on this grassland range community, however, was the remarkable diversity of forbs in combination with dominance of decreaser tallgrass species. Forb species included black-eyed susan (Rudbeckia hirta), the conspicuous yellow-flower composite, prairie gayfeather or prairie blazing star (Liatris pycnostachya), several goldenrod (Solidago) species, Baldwin ironweed (Vernonia baldwinii), roundhead lespedeza (Lespedeza capitata), compass plant (Silphium laciniatum), and common horsemint or wild bergamot (Monarda fistulosa). And all of these listed grass and forb species in the frame of this one photograph!

Prairie State Park, Barton County, Missouri. July. Estival aspect. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). Mesic Prairie. SRM 710 (Bluestem Prairie) and SRM 802 (Missouri Prairie). Central Irregular Plains- Cherokee Plains Ecoregion, 40d (Chapman et al., 2002).

74. Heartland Garden of Eden: quintessential tallgrass prairie range- Here was tallgrass prairie in its ultimate expression as the climatic climax of its zone. Mesic Prairie community with big bluestem (just reaching anthesis stage; many sexually reproductive shoots still in the boot) the dominant and Indiangrass the associate species. This outstanding example was from a gradual mid-slope upland site in close proximity to the species-rich example shown in the immediately preceding photograph. Both of these "photo-plots" were in the Cherokee Prairie Region. This was in the Cherokee Lowlands portion of the Osage Plains section of the Central Lowlands physiographic province (Fenneman, 1938, ps. 455, 605-630 [esp. 612-613]).

At one time this general area was known for its world-class prairie hay. Several towns and counties in southwest Missouri and southeast kansas are the self-proclaimed "Prairie Hay Caipital of the World". Obviously there is no "one right answer", but the assertion of Minden Mines, Missouri for this coveted title bears special merit.

Prairie State Park, Barton County, Missouri. July. Estival aspect. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). Mesic Prairie community. SRM 710 (Bluestem Prairie) and SRM 802 (Missouri Prairie). Central Irregular Plains- Cherokee Plains Ecoregion, 40d (Chapman et al., 2002).

Diversity in a prairie Garden of Eden- A rich assembly of native prairie plant species on the virgin sod of an upland range site in the Osage Plains of western Missouri. Tallgrass prairie vegetation seen in this "photo-quadrant" was dominated by big bluestem with switchgrass the associate species. This was a remarkable array of composite forbs ranging from the perennial decreaser, Maximillian sunflower down to the annual invader, giant ragweed (Ambrosia trifida), with common ragweed (Artemisia artemisifolia), a native perennial invader, in between so to speak. The mint family (Labiateae) was represented by slender mountain mint (Pycanthemum tenuifolium), a native perennial found on climax tallgrass vegetation, while the carrot family (Umbelliferrae) had as its representative the Eurasian, naturalized, biennial forb known as wild carrot or Queen Anne's lace (Daucus carrota).

Relict tract, Missouri Prairie Foundation, Vernon County, Missouri. July. Estival aspect. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). Mesic Prairie community. SRM 710 (Bluestem Prairie) and SRM 802 (Missouri Prairie). Bluestem "Tall-Grass" Series 142.11 of Plains Grassland 142.1 of Cold Temperate Grassland 142 of Brown et al. (1998, p. 40). Dry-Mesic Limestone Prairie (Nelson, 2010, ps. 277-281). Central Irregular Plains- Cherokee Plains Ecoregion, 40d (Chapman et al., 2002).

75. The healing sward of tallgrass prairie- This view of the sward — the aboveground portion of the vegetation of marsh, meadow, or grassland; the herbaceous surface of a grassland or other graminoid-dominated community —of tallgrass prairie shows the characteristic combination of cespitose (clumped or tufted ) grasses or bunchgrasses that reproduce asexually (vegetatively) with tillers (vertical or intravaginated shoots) and the sod-forming grasses that reproduced vegetatively with rhizomes and/or stolons (extravaginated or horizonal shoots belowground and aboveground, respectively). Some grasses such as big bluestem, which is the dominant species here, and Indiangrass and switchgrass, the main associates in this community, have both tillers and extravaginated shoots like rhizomes (in all three of these species). Other grasses are strictly bunchgrasses with shoots limited to tillers like prairie dropseed the third associate seen in this view.

Note to beginning students: shoot is the above ground portion of grass and includes culm (grass stem), leaves, and inflorescence. Tiller is not a synonym for shoot except when the shoot is tiller, a vertical intravaginated shoot. Tiller is not the generic for shoot. It is incorrect to speak or write of “tiller dynamics” when shoots being discussed include stolons (“runners”) or rhizomes. Then it is shoot dynamics. Freshman Agronomy 105.

This virgin sod (sod is the combination of roots and the soil they hold or bind; it is the belowground strata of grassland, marsh, or meadow and usually connotes the first few inches of aboveground vegetation) shows the perfect soil- protecting capacity of shoot and root of grass. It also illustrates the erosion-healing feature of grass. Note the “haired-over” old gully in the background. Senator Ingalls of Kansas had it right: “Grass is the forgiveness of nature…”

There are various species of forbs in this Excellent range condition sward but as typical for pristine prairie, the great bulk of biomass (weight of living orgamisms or, more precisely, the live weight of organisms) is contributed by the grasses, and often just two up to maybe five or six species (four in the example seen here). This is mid-estival aspect with the grasses in the boot stage (the phenological stage in which the inflorescence is is still enclosed or enveloped by sheath of uppermost leaf). The short- shoot grasses (those which do not elevate their apical meristem until later in the growing season) like big bluestem are just beginning to elongate their culms. July.

Based on herbage yield, palatabability and nutritive value of growing forage, resilence (recovery) from disturbance, soil formation capability, habitat for number of wildlife species, and contributions to Indian and cowboy culture, this is perhaps the greatest natural pasture in North America if not on Earth. God's own cow country; a rangeman's Garden of Eden.

Tallgrass Prairie Preserve (the former Chapman-Barnard Ranch) Osage County, Oklahoma. FRES No. 39 (Prairie Ecosystem), K-66 (Bluestem Prairie), SRM 710 (Bluestem Prairie). Flint Hills Ecoregion, 28a (Woods et al., 2005).

76a. The ultimate in heartland grasslands- Tallgrass-true prairie expression of climax grassland in the interior of the humid zone of North America. A local prairie dropseed consociation in a predominately big bluestem-switchgrass mesic tallgrass prairie. Othe major species within this small-scale mosaic of grassland communities (a patchwork of small areas of several contiguous range sites) were porcupine grass (Stipa spartea) and various species of caric sedge (Carex spp.). The interpretation and distinction between true prairie and tallgrass prairie as elaborated by Weaver and Clements (1938, ps. 458-460, 516, 518-521) was described in the chapter, True Prairie, herein.

The vegetation of this pristine prairie, one of the jewels of the preserved prairies in Missouri, seemed to be a transition-like expression of grassland vegetation between tallgrass prairie and true prairie range types. However, this "hybrid-like" combination grassland community could not be readily regarded as an ecotone (as ecotonal) or a true transition between the two distinctive types give that they were not conterminous. Rather, there were just scattered consociations of prairie dropseed in a big bluestem-dominated tallgrass prairie. In the context of Landscape Ecology there were patches of prairie dropseed in a matrix of big bluestem. Populations of upland switchgrass on mima mounds (often populating them almost exclusively) could also be regarded as patches in a big bluestem matrix.

Diamond Grove Prairie, Newton County, Missouri. July. Prebloom stage of prairie dropseed; pre-shoot elongation stage of big bluestem. Estival aspect. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). Mesic Prairie community. SRM 710 (Bluestem Prairie) and SRM 802 (Missouri Prairie). Soil Survey of Newton County, Missouri (Soil Conservation Service, 1992) did not include range sites ("Go figure"). Soils here were Hoberg-Keeno (silt loams), Gerald (silt loam), and Credon (very cherty loam) associations. Ozark Highlands- Springfield Plateau Ecoregion, 39a (Chapman et al., 2002).

76b. Another grassland community in the ultimate of heartland grassland- A local range plant community of sloughgrass (= prairie cordgrass) and eastern gamagrass on the overall big bluestem-dominated tallgrass prairie introduced in the preceding photograph. This component part of tallgrass prairie vegetation was another form of plant community patch, a much more mesic one, in the matrix of the big bluestem-dominated grassland.

Diamond Grove Prairie, Newton County, Missouri. July. Grain-ripe stage of eastern gamagrass; pre-boot stage of sloughgrass (= prairie cordgrass). Estival aspect. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). Mesic Prairie community. SRM 710 (Bluestem Prairie) and SRM 802 (Missouri Prairie). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Chapman et al., 2002).

76c. More views of the grassy heartland- Three landscape-scale views of a dry-mesic chert prairie with features of both the tallgrass prairie and true prairie associations (Clements, 1920, p. 121-134; Weaver and Clements, 1938, ps. 518-523). Co-dominant species were big bluestem and switchgrass (an upland ecotype) while prairie dropseed was the associate species over much of this ozark (Springfield) Plateau grassland. With these relative proportions along with the drier habitat of this grassland this climax vegetation had features of both Clementsian associations.

Another major grass species was silky wildrye (Elymus villosus). This was one of the few major grass species on this prairie that was a cool-seaon species (subfamily, Festucoideae; tribe, Hordeae or Tritaceae). The vast majority of the grass species (and almost all the grass cover) on this heartland prairie were warm-season grasses of the Panicoideae and Eragrostoideae (the panicoid or eragrostoid subfamilies).

The major grasses like big bluestem, switchgrass, and prairie dropseed grew in local populations across this grassland landscape so that there were spots of consociations of these grass species. This is often a characteristic pattern of vegetation on climax grasslands. Clements (1920) recognized the phenomenon of a single species dominating local--often large local--"patches" which is why he coined the term (Clements was good at [or bad about] coining terms), consociation. In the second and third of the three slides presented here most of the grassland herbage was that of big bluestem. These were two separate consociations of big bluestem.

There were two important (defining) species of grasslike plants in the virgin sod of this climax range plant community: 1) Mead's caric-sedge (Carex meadii) and 2) grassleaf rush (Juncus marginatus). Two of the major families of grasslike plants were thereby represented: 1) Cyperaceae and 2) Juncaecae. Most of the prairie forbs in this grassland had completed blooming and fruit production. A major prairie forb in the three views shared here that was still in flower was finger poppymallow or finger winecups (Callirhoe digitata). The most common forb in these three views was foxglove beardtongue (Penstemon digitalis) which was in the ripening-fruit stage.

Visible in these three photographs were mottes or small patches of hardwood species. Regular occurrence of natural prairie fires kept these to a minimum in the pre-whiteman era when American Indians did a far superior job of Range Management than the whiteman has achieved. An example of such hardwood patches was shown and discussed in detail below. Sadly, much of the land surrounding this pristine prairie developed into disturbance climaxes (another of Clements' terms--and, typically, a useful term) of oak-hickory forest under influence of underburning and overgrazing. Use of this prairie for the production of prairie hay for over 150 years was the only thing that kept it as such a crown jewel of the Show Me State that was once probably one-third covered by prairie.

Schroeder (1982) closely examined Missouri land surveyor reports and concluded that presettlement Missouri was 27% coverd by prairie. This author suspects that the proportion was slightly higher than that. Anyway this prairie in the western Ozark Plateau was preserved thanks to self-less prairie disciples. Thank you.

Technical note: an Epson Perfection 700 scanner over-exposed all three of these Provia 100F slides creating a "sparkle" appearance to grass shoots. Compare the "frosty" or "sparkled" appearance or texture of thee three slides to the two slides in the two immediately slide/caption units whcih were scanned with a Hewlett-Packard scanner which required hand-scanning of each slide. This was not a controlled experiment. The two immediately preceding slides were Kodachrome 65 film, and the HP scanner always made the tones a bland or muted "off-color". Kodachrome did not have the bluish or greenish cast produced by the Hewlet Packard scanner. Upshot: there are no high-quality slide scanners. Ostensibly, they do not exist. One has to make do: this photographer is not about to give up slides, hard copy images, for digital vapors stored on a compact disk, portable hard-drive, or "the cloud".

Diamond Grove Prairie, Newton County, Missouri. Early August; mid- estival aspect, early bloom stage of tallgrass species. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). Dry-mesic chert prairie (Nelson, 1987). SRM 710 (Bluestem Prairie) and SRM 802 (Missouri Prairie). Bluestem "Tall-Grass" Series, 142.11, of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40).Ozark Highlands- Springfield Plateau Ecoregion, 39a (Chapman et al., 2002).

77a. About droughted out- Another part of the same tallgrass prairie in southwestern Missouri as was featured in preceding slides. In this part of the dry-mesic chert prairie the effects of Severe Drought (Palmer Drought Index) were more evident than in parts of this prairie shown in the three slides of the immediately preceding set. There was a greater proportion (cover) of Meads caric-sedge which had completed its annual growth cycle so that was a major factor that influenced the appearqnce of this "photosample" of the prairie. Mead's caric-sedge and prairie dropseed were co-associate species in the grassland vegetation seen here. Big bluestem and switchgrass (an upland ecotype) were co-dominants of this climax grassland community.

The high proportion of prairie dropseed along with the two more mesic members of the Four Horsemen of the Prairies tallgrass species (little bluestem and Indiangrass being the more xeric or less mesic of the four major tallgrasses) indicated the affinity of this Ozark (Springfield) Plateau grassland with both the tallgrass prairie and the true prairie associations (Clements, 1920, p. 121-134; Weaver and Clements, 1938, ps. 518-523).

77b. Less drought-stricken- This interesting set of prairie plants wass photographed at the same time as the immediately preceding slide, but this microhabitat (=microsite= microenvironment) had not suffered drought to the extent or degree as the tallgrass prairie vegetation presented in the preceding slide. This demonstrated the local variation possible in range ecosystems. "Deeper" soil, soil with greater water-holding capacity, is always a "prime suspect" factor in such circumstances.

Tallgrass prairie species in this "photoplot" included the featured false or whhite wild indigo (Baptsia leucantha) loaded with legumes as well as a stray wild plum (Prunus sp.?) along with co-dominants big bluestem and switchgrass (an upland ecotype), and shining or smooth sumac (Rhus glabra).

78. Mima mound topography- Mima mounds are frequently characteristic of the virgin sod of tallgrass and true prairies. These "prairie pimples", "Indian mounds", or "meadow biscuits" have long intrigued prairie hay makers as well as ecologists. Laymen and scientists alike have advanced pet explanations for this unique example of micro-topography. This pattern of microrelief (Soil Survey Division Staff, 1993, ps. 69-70) is similar to the gilgai phenomenon, but the cause(s) are not as obvious as in the shrink-swell of gilgai clay soils. Knight (1994, ps. 125-130) discussed these "puzzling landscape features" in the Laramie Basin as to their origin and possible effect on vegetation. While this microrelief has received quite a bit of study (Knight provided numerous citations) findings are still inconclusive though mound vs. intermound vegetation is characteristically different.. That is the situation for this excellent condition prairie hay meadow in the Missouri Ozarks. Switchgrass and prairie cordgrass occupy the mounds while big bluestem and prairie dropseed dominate intermond soil.

This is similar to the Clementsian concept of postclimax vegetation on deep sand sites (eg. tallgrass vegetation on sandhills in a semiarid mixed prairie region). Grasses of the mounds are clearly the more mesic species. The maroon flowers are those of tall winecup (Callirhoe digitata) which grows only on the mounds. This illustrates the concept of an indicator plant. Mima mounds are a textbook example of microhabitat or –environment within a range site.

Diamond Grove Prairie, Newton County, Missouri, July. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). Dry-mesic chert prairie (Nelson, 1987). SRM 710 (Bluestem Prairie) and SRM 802 (Missouri Prairie). Bluestem "Tall-Grass" Series, 142.11, of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40).Ozark Highlands- Springfield Plateau Ecoregion, 39a (Chapman et al., 2002).

79. Freshly mowed mima mound- A mima mound in a meadow recently mowed for prairie hay immediately adjacent to the mound in the immediately preceding photograph. This on-the-mound photograph presented a more detailed view at sward level of this microtopography that is a characteristic feature of tallgrass prairie grasslands in the Ozark Plateau and Cherokee Prairie.

80. Tallgrass prairie in Ozark Plateau- Four Horsemen tallgrass prairie with big bluestem and switchgrass co-dominants and prairie dropseed the associate species (Indiangrass and little bluestem were fourth and fitth most abundant species, respectively). Two photographs of broad scale-views showing physiogonomy of range plant community and physiography of land, including microrelief of mima mounds. Range vegetation in first slide included a lot of tall nut-rush or whip nut-rush (Scleria triglomerata) and prairie gayfeather or prairie blazingstar (Liatris pycnostachia). The second slide showed a portion of range vegetation with many more prairie forbs including wingstem or yellow crownbeard (Verbesina helianthoides), finger poppymallow or winecups (Callirhoe digitata), smooth penstemon or foxglove beardtongue (Penstemon digitalis), and rosinweed (Silphium integrifolium).There was also a lot of Virginia wildrye (Elymus virginicus) in vegetation seen in second photograph along with quite a bit of bolting (rapid development of sexual or flower shoots) in big bluestem.

This was a dry-mesic chert prairie (Nelson, 1987; 2010, ps. 281-284).

Range students correctly associate tallgrass prairie with such well-known grassland areas as the Flint Hills or Nebraska Sandhills, but beginning students commonly think only of forests with regared to natural vegetation of the the Ozark Mountains. In actuality much of the native vegetation of the Ozark Plateau is tallgrass prairie and tallgrass-oak/hickory savanna. This short section of Ozark Plateau tallgrass prairie was introduced to overcome that incorrect perception.

Diamond Grove Prairie, Newton County, Missouri. Late June, early estival aspect. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). Dry-mesic chert prairie (Nelson, 1987). SRM 710 (Bluestem Prairie) and SRM 802 (Missouri Prairie). Bluestem "Tall-Grass" Series, 142.11, of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40).Ozark Highlands- Springfield Plateau Ecoregion, 39a (Chapman et al., 2002).

81. Details of an Ozark tallgrass prairie- Structure and botanical composition of a Four Horsemen tallgrass prairie in which prairie dropseed was also a dominant--even first dominant species. The lighter-green, cespitose plants with downward, sprawling leaves (and parted tussock) crown were prairie dropseed. White forbs in first slide were slender or narrowleaf mountain mint (Pycanthemum tenuifolium). Long, dead grass shoots in first slide were those of big bluestem. Plants of the prominent yellow composites in second slide were black-eyed Susan (Rudbeckia hirta) and the large-leafed forb in midground were plants of prairie tea (Croton monanthogynus). Also in the second slide were numerous shoots of Junegrass.

82. Tallgrass prairie distinguished by abundance of prairie dropseed- Details of sward of Four Horsemen plus prairie dropseed on an Ozark prairie. The three most abundant plant species in this virgin tallgrass prairie were big bluestem, switchgrass, and prairie dropseed. Switchgrass was the dominant on mima mounds while big bluestem was the dominant on microsites of mid-mesic conditions and prairie dropseed was the dominant on less mesic microhabitats. The large, cespitose plants with drooping (ascending and then downward facing) leaves in center field of both of these "photoquadrants" were prairie dropseed.

Prairie dropseed was the distinctive species throughout much of this grassland being either dominant or associate species over many microenvironments. Weaver and Clements (1938, p. 518) explained that prairie dropseed was one of three characteristic dominants of the true (in contrast to tallgrass) prairie. Presence of prairie dropseed on this obvious tallgrass prairie demonstrated the close botanical affinity and spatial proximity of these two major types or associations of mesic midcontinental grasslands.

Other conspicuous plant species in these two slides were slender mountain mint (bright, white inflorescences), black-eyed Susan (yellow composites), and tall or whip nut-sedge (immediate center foreground of second slide).

83. A unique one featured- This "photoquadrant" of the sward of a tallgrass prairie in the Ozark Plateau highlighted the annual member of Gentianaceae (gential family) commonly known as rose pink or rose gentian (Sabata angularis). This is one of the few native annual forbs that is found in tallgrass prairie in "mint condition" (ie. as a member of the virgin sward). As with all plant species, this plant was described in standard flora of this region including Stayermark (1963) and McGregor et al. (1986). When in blooom this forb really "sets off" the turf of an already spectacular prairie.

Diamond Grove Prairie, Newton County, Missouri. Late June.

84. Surrounded by woods- Landscape of a tallgrass prairie in the Ozark Plateau of southwest Missouri showing margin of this pristine grassland surrounded by what--except for haying and prescribed burning--was encroaching oak-hickory forest. Interpretation of the natural vegetation of the Ozark Plateau requires careful analysis in context of vegetation development at both geological and successional temporal scales. The vast populace of unenlightened viewers of Ozark vegetation (including otherwise well-educaed foresters) commonly envision the Ozarks as forests of hardwood (primarily oak-hickory) or hardwood-pine, but this is a misconception or, at very least, an only partly true sterotype. In reality much of the tree-dominated natural vegetation of the Ozark Plateau consisted of 1) open forests with understories of tallgrass prairie grasses, especially big bluestem, 2) savannas of oaks and hickories with lush tallgrass (or tallgrass and shrubs) understories, 3) grasslands (primarily tallgrass prairies such as the one shown here), and 4) glades of tallgrass and forb species. Development of oak-hickory or oak-hickory-pine forests with closed canopies and understories largely devoid of herbaceous species and instead comprised primarily woody layer(s) was a vegetational phenomenon attributable largely to fire exclusion by the actions--direact or indirect--of white man (eg. farming;, roadbuilding; overgrazing; and planned, Smoky bear fire exclusion).

This afforestation (establishment [in this case by human action] of a forest on land where the preceding vegetation was not forest [Helms, 1998]) of the Ozark Plateau was well-documented and plainly described by numerous workers. Readers were referred to the classic works of Steyermark (1940, 1959) along with some of the other studeis described above. Similarily, much of Missouri as well as adjoining portions of Illinois and Iowa were almost exclusively grasslands of true and tallgrass types (Steyermark, 1963, ps, xxii-xxiv). Tallgrass prairie like this one in the western border of the Springfield Plateau section of the Ozark Plateau was typical in physiogonomy, species composition, structure, and function to those of the general and vast Prairie Region.

Diamond Grove Prairie featured in this section had been kept free of woody invaders for decades through haying (harvest for prairie hay) long before it was purchased in 1972 through generosity of Miss Katherine Ordway and incorporated into the outstanding system of prairies administered by one agency or another of the Show Me State of Missouri. (A photograph of part of the recently hayed Diamond Grove Prairie was presented above.) Beginning several years before time of this and subsequent photographs haying operations ceased on Diamond Grove. Invasion of the former Diamond Grove hay meadow by native woody species bcame obvious within two or three years after mowing for prairie hay. Prescripton burning of Diamond Grove Prairie was initiated by Missouri Department of Conservation when it became obvious that maintenance of this virgin grassland was best--at least, most naturally--achieved by proper use prescribed fire.

Forested land seen on the margin of Diamond Grove Prairie (distant background of both slides) was private property that had been overgrazed by cattle for decades and, most likely, farmed prior to that. The virgin grassland of Diamond Grove Prairie had been preserved only because it had been set aside by various landowners for production of prairie hay. Once mowing for hay production was curtained Missouri Department of Conservation invasion of the tallgrass prairie by native species of woody plants began. The agency was forced to use prescribed burning or, alternatively, watch the virgin prairie become an unnatural forest as a result of the artificial exclusion of prairie fires. Fortunately, the agency spurned Smokey Bear and wisely applied proper prairie use through prescription burning.

Native vegetation presented in these two photographs consisted primarily of five grass species: big bluestem, switchgrass, prairie dropseed, Indiangrass, and little bluestem (in that relative, general order). Switchgrass was the dominant species on mima mounds such as the one featured in the second of these two slides. Major forbs included slender mountain mint, smooth or foxglove beardtongue, prairie blazingstar or prairie gayfeather, leadplant, and black-eyed susan. Woody (= invader) species persent as seedlings to small saplings included blackjack oak (Quercus marlandica), post oak (Q. stellata), blackberry or briar (Rubus spp.), persimmon (Diospyros virginiana), and sassafras (Sassafras albidum). These woody invaders were "kept at bay" only through prescribed burning (see immediately below).

85. Woody invaders of an Ozark tallgrass prairie- Within three months of prescribed burning there were resprouts of blackjack oak (first photograph) and post oak (second photograph) in tallgrass prairie vegetation of Diamond Grove Prairie. Also visible in the second photograph were plants of persimmon and wild plum (Prunus sp.) Once hay-making operations were curtailed (several years prior to time of photographs), and in combination with minimal (only incidental) grazing by white-tailed deer and smaller herbivores, invasion of the virgin tallgrass sod by woody plant species began. Adjacent oak-hickory forest on the perimeter of Diamond Grove Prairie (contiguous or conterminous forest vegetation) as well as the soil seed bank served as sources of propagules or disseminules for this woody invasion.

Plants of the two oak species, persimmon, and wild plum had established one or two years prior to this year's (current spring) prescribed burn. Once established, most of such woody plants can be kept at immature stages of development only through recurrent prairie fires.

Black-eyed Susan and slender mountain mint were the conspisuous forbs in the first slide. Junegrass was noticable in the second slide.

Diamond Grove Prairie, Newton County, Missouri. Late June, early estival aspect. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). Dry-mesic chert prairie (Nelson, 1987). SRM 710 (Bluestem Prairie) and SRM 802 (Missouri Prairie). Bluestem "Tall-Grass" Series, 142.11, of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Chapman et al., 2002).

86. Haymaker's mark- A foolproof sigh that a prairie is being (or has, in the past, been) used as a hay meadow is presence of patches of woody vegetation scattered within the grassland plant community. It is true that annual mowing for hay insures that there are no aboveground parts (shoots) of woody plants older than one year in mowed areas; however, mowing machine operators give a proverbial "wide berth" to any woody material large enough to damage their equipment (eg. break sections in sicklebar mowers, dull blades of disk mowers). Leaving unmowed parts of hay meadows results in patches or strips of shrubs, tree sprouts, etc. The result is ever-expanding patches of brush. In fact, year after year these areas of woody plants increase exponentially as their avoided perimeters with woody plants too large to be mowed encroach outward into herbaceous vegetation in roughly circular patterns. The geometric boundaries of brush patches that grow outward each year extend at exponential rates of invasion. This is a ripple-like or wave-resembling encroachment of brush into what was previously grassland. Over time successively greater areas of prairie are invaded to eventually become woodland or forest.

The only mechanical form for control of brush (reduction in aboveground cover of noxious woody plants) in prairies, such as those used for hay meadows, is cutting or digging by hand or, more realistically, shredding (breaking or twisting off of shoots by rotary brush cutters). Over large areas such mechanical control is expensive and not always cost-effective. Broadcast chemical control such as aerial application of herbicide is out of the question because herbicides that kill woody docots will also kill desirable forbs such as native legumes and palatable composites. Chemical control will not permit preservation of the natural vegetation. Fire is the only viable alternative in such instances. That is why the Missouri Department of Conservation was compelled--eventually--to use prescribed burning to maintain Diamond Grove Prairie.

This Ozark Plateau prairie needed more flames a lot sooner, but "better late than never". At this late point in time (stage of brush invasion) some of the trees in Diamond Grove Prairie were already too large to be topkilled (and root kill was already at the point of being "highly unlikely") by prescription burning. The only feasible approach for brush control at this stage of woody invasion (as indicated by size of established trees) was either spot application of herbicides or physical removal of trees with equipment like bulldozer, grubber, or track hoe. "An ounce of prevention is worth a pound of cure". The commendable use of prescribed fire was already "too little too late".

The first slide of this set presented an overall view of a patch of burnt brush that included blackjack oak, black oak (Quercus velutina), post oak, sassafras, persimmon, smooth sumac (Rhus glabra), and wild plum (Prunus sp.) in that approximate order of abundance and cover. The second slide was a close-up view of fire effects in this brush patch. Smooth sumac presented a special situation of brush control because the various Rhus species greatly and rapidly increase in cover and density (much like big bluestem) under late winter or spring burning. Expansion of smooth sumac into previously sumac-free grassland as a direct response to the three-month earlier prescribed fire was obvious in the second slide. Rhus species are some of the most fire-adapted--in fact, fire-dependent--of all prairie shrubs. This lesson was showed in greater detail in the next slide-caption set.

The largest and second largest trees in middle of this brush patch were black oak, one of the most widespread dominants of oak-hickory forests throughout the western (Springfield) portion of the Ozark Plateau. This aggregation of woody plants was already becoming the "nucleus" of an Ozark forest. The climax vegetation of the Ozark Region was a mosaic of oak-hickory forests; grasslands (mostly tallgrass prairies and isolated glades over shallower, rockier soils); and savannahs of oak and hickory trees, scattered shrubs, and understories dominated by tallgrass species. Recurrent fire was a major "landscape architect" responsible for this patchwork of native plant communities. (This phenomenon was described in greater depth in the next photocaption.)

87. Too much fuel; too little fire- Two successively closer views of the brush patch introduced immediately above that developed on a tallgrass prairie hay meadow in the Springfield (western Ozark) Plateau. This woody vegetation developed due to a combination of fire cessation (fire suppression) and lack of mowing. Hay mowers avoided groups of shrubs and small trees resulting in patches of woody vegetation that invaded the virgin grassland at expontential rates (ie. rates of woody invasion increased at annually increasing geometric rates).

Woody species were primarily blackjack oak, black oak, post oak, sassafras, persimmon, smooth sumac, and wild plum (in roughly that order of relative cover). The largest plants, including the conspicuous tallest plant, in these two slides was blackjack oak. This patch of woody plants was already on its way to becoming an oak-dominated forest!

The brush patch presented in these two and the two preceding slides had been treated with prescribed fire about three months prior to time of photographs. Obviously many iof the woody plants were not even topkilled and apparently none of these brush plants had been rootkilled. Instead prescribed burning resulted in proliferation of the highly fire-adapted smooth sumac (preceding two slides) and profuse sprouting from rootcrowns and rootstocks (woody rhizomes) of sassafras. Resprouting of sassafras was featured in foreground of the two photographs.

Repeated burning (on, say, a recurrent cycle of one to every three years) would likely reduce (eventually rootkill) many of these woody plants, but they were certainly well-established by the time the Missouri Department of Conservation belatedly employed prescribed fire, an indespensible tool for maintenance of such climax grasslands in this humid zone. In absence of fire--a natural outcome of a continental climate with profuse lightening strikes--climax tallgrass prairie would develop into postclimax oak-hickory forest or, at very least, become a degraded grassland invaded by woody species that are less fire-tolerant than climax herbaceous species.

Under humid continental climate, tallgrass prairie is more likely a pyric climax (= a fire type) than a climatic climax. This is, however, a play on words used to designate (and emphasize) a so-called fire climax. Properly (or more accurately) stated , the fire component (outcome) of climate determines the prevailing potential natural vegetation. Fire is merely the resultant phenomenon of humid precipitation that produces abundant, readily flamable fuel which is easily ignited by prolific strikes of lightening and rapidly spread by usually amble wind speed. Recurrent fire is the culminating feature of climate that is largely responsible for prevention of establishment (or elimination and drastic reductions) of plant species that have aboveground perennating parts. Herbaceous plants which have almost no perennial organs aboveground are favored over woody plant species under recurrent fire, a final feature or facet of climate. Fire-conducive climates (regions having frequent natural fire as part of their prevailing climate) have herbaceous rather than woody plant communities as the climax vegetation. Hence, the term of fire climax to lay stress upon the final climatic feature responsible for climax grasslands.

This is more the case when fire burns herbaceous plants during dormancy. Even when fire burns herbaceous plants during growing their growing season these species with their annual aboveground organs are less severely damaged than woody plants that have years of accumulated aboveground growth.

88. Season's end; herbage at its peak- Appearance of largely ungrazed biomass at peak (or near it) Early Auguststanding crop on a tallgrass prairie in the Springfield Plateau of southwest Missouri. Major grassland range species were big bluestem, the overwhelming dominant; switchgrass and Indiangrass, the associate species; and prairie dropseed and Mead's caric sedge (Carex meadii) as next "in line" based on estimated cover, plant density, and biomass. Little bluestem was "conspicuous by its absence". Forbs were very limited with the main two species within view of these two "photoplots" being great or giant ironweed (Vernonia crinita) and blue or swamp vervain (Verbena hastata).

Woody patche in distant background were colonies of winged or shining sumac (Rhus copallina), a species that had increased under recent prescribed burning (revisit slide/caption sets immediately above).

This prairie had been used as a prairie hay meadow for decades (probably upwards of a cetury as matter of fact) and had not been subject to livestock grazing for that hay meadow period. This prairie was grazed/browsed by white-tailed deer (Odocoileus virginianus) and smaller herbivores, including a myriad of insect species.

Diamond Grove Prairie, Newton County, Missouri. Early August, estival aspect. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). Dry-mesic chert prairie (Nelson, 1987). SRM 710 (Bluestem Prairie) and SRM 802 (Missouri Prairie). Bluestem "Tall-Grass" Series, 142.11, of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40).Ozark Highlands- Springfield Plateau Ecoregion, 39a (Chapman et al., 2002).

89. The glory of tallgrass prairie in the spring- A treasured tallgrass hay meadow dominated by big bluestem with Indiangrass as the associate species as seen in vernal aspect. The conspicuous prairie forb was American cowslip or shooting star (Dodecatheon meadia). "[E]ven Solomon in all his glory was not arrayed as one of thes" (Matthew 6:29; Luke 12:27) -- and we've got a whole meadow full.

Ottawa County, Oklahoma. April. Peak bloom in cowslip. FRES No.39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 710 (Bluestem Prairie). Plains Grassland 142.1, Bluestem "Tall-Grass" Series, 142.11 of Brown et al. (1998, p. 40). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al. (2005).

Prairie Cordgrass Wet Prairie

Wet prairie dominated by prairie cordgrass (Spartina pectinata) was once (before the tile-laying, sod-busting farmer) a major form of tallgrass prairie.The whiteman converted almost all of this to farm fields. Wetlands--especially those with rich soil suitable for tillage and production of field crops--were viewed in the same way as virgin forests. They were something to be converted as quickly and efficiently as possible to fertile farms of the yeoman farmer. Thrown in fear of the "ague" (malaria) and the grassland stage was set for destruction of wet prairie so as to convert these hydric habitats to mesic farm land.

Today even with a conversion of human perspective to regarding wetlands as some of the most unique and treasured of all natural habitats, few relicts of wet prairie remain. Fortunately far-sighted conservationists and prairie lovers managed to save some relicts of prairie cordgrass or "ripgut", as cordgrass is known by locals, wet prairie in the Osage Plains of western Missouri. Some of hese priceless tracts of virgin "ripgut" wet prairie were and sre still continuing to be purchased and preserved by selfless grasslanders, who in cherishing prairies, are leaving these gifts as a natural legacy to the present and future generations of prairiemen.

The following images were taken The Nature Conservancy's Marmaton River Bottoms Wet Prairie Preserve and the adjoining private property of

90. Relict of wet prairie- The virgin sod of a consociation of prairie cordgrass was the climax grassland community of this wet prairie on the floodplain of Marmoton River in the Osage Plains of western Missouri. Trees in the distant background were along the closer terraaces of Marmoton River. The first slide featured a local small colony of buttonbush (Cephalanthus occidentalis), climax (decreaser) shrub of wet to aquatic environments.

The shoots of prairie cordgrass in these and the following views of a prairie cordgrass consociation on floodplain of Marmaton river were in late pre-bloom to early bloom stage of phenology.

Local folks in this area know prairie cordgrass as "ripgut".

The Nature Conservancy's Marmaton River Bottoms Wet Prairie Preserve, Vernon County, Missouri. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie), but this is a variant of that so Kuchler unit not really descriptive. Likewise, SRM 802 (Missouri Prairie) is not adequate because this is a wet prairie or prairie cordgrass variant thereof. Cold Temperate Grassland 142, Plains Grassland 142.1, Bluestem "Tall-Grass" series 142.11, prairie cordgrass variant or sub-unit of (Brown et all, 1998, p. 40). Range site: Clay Lowland (Soil Conservation Service, 1981). Central Irregular Plains- Cherokee Plains ecoregion 40d (Chapman et al., 2002).

91. Beauty in sameness- Virgin stand of prairie cordgrass, a consociation of this species, on wet prairie in the Osage Plains of western Missouri. This climax grassland vegetation developed on a heavy clay soil on the floodplain of Marmoton River. Local farmers refer to such dense, wet clay as "gumbo" or "gumbo" clay. Prairie cordgrass vegetation can be handily thought of as an edaphic climax in the polyclimax theory of Arthur Tansley. In the monoclimax theory of Clements prairie cordgrass on 'gumbo clay" would be post climax.

Either way the sameness or homogenous composition of this marvelous wet grassland was the farthest thing from a vegetational monotony. To prairiemen, the constancy, with variations (some quite slight), of the great grasslands is one of the endearing features for which they love the prairie. To a prairieman like your author the vast expanse of sameness, a natural grassland as constant and consistent in its composition as a farmer's monocultured field, is awe-inspiring, inspirational, and the height of aesthetic pleasure.

Three views of sheer grandure shared with some who have never experienced joy.

92. Two more inspiring views- A consociation of prairie cordgrass on the Marmaton River floodplain with an nearly 100 percent stand of prairie cordgrass in the first slide with trees (primarily pecan with some bur oak), closer to Marmaton River. In the second slide a local stand of Pennsylvania smartweed (Polygonum pensylvanicum) in the foreground.

93. Haying God's own river bottom- The shorter shoots of prairie cordgrass at left is regrowth following hay-making two or three weeks earlier while the unmowed shoots at right indicated height of prairie cordgrass awaiting the mowing machine. Trees in the background were on the closer floodplain of the Marmoton River.

Location note: a bottomland forest (or, in areas where trees were more widely spaced, woodland) that was dominated by (was a consociation of) native pecan (Carya illinoiensis) was contiguous with (immediately adjacent to) the prairie cordgrass wet prairie on the floodplaind of Marmaton River (Vernon County Missouri) presented immediately above. Treatment of that pecan-dominated forest range with an understorey of Canada wildrye and broadleaf woodoats (Uniola latifolia) was in placed in two chapters of this publication: Miscellaneous Forest Types-I (variously throughout that chapter--keep looking).

94. Prairie Cordgrass (Spartina pectinata)-dominated community of tallgrass prairie- Clay bottomland range site. Note wild indigo (Amorpha fruticosa).Anderson County, Kansas. Estival aspect, July. FRES No. 39 (Prairie Ecosystem). Bottomland variant of K-66 (Bluestem Prairie). SRM 710 (Bluestem Prairie) variant or SRM 601 (Bluestem Prairie) variant. Central Irregular Plains- Osage Cuestas Ecoregion, 40d (Chapman et al., 2001).

95. Prairie cordgrass or sloughgrass (Spartina pectinata) growing conspicuously in a slough on tallgrass prairie- This species and the local low spot appeared quite pronounced in a dry year. Spartina was derived from the Greek, spartinae, meaning "cords" probably in reference to the tough leaves (Hitchcock and Chase, 1951, p. 508), but many know this particular species more by its major habitat of small damp draws called sloughs. Ecologists like John Weaver (1954, ps. 31-33) more commonly called this most water-requiring of all the major tallgrass species sloughgrass or tall marshgrass. Weaver (1954, p. 33) noted that sloughgrass was "the last stage in the succession from wet land or water to climax prairie" and that it formerly formed essentially single species-stands (consociation in the Clements-Weaver unit of climax vegetation) over hundreds of square miles of bottomland along watercourses such as the Mississippi and Missouri Rivers. Almost all of this riparian range (and that of the draws on upland prairies like the one shown here) was drained, plowed, and planted to row crops, especially corn. This relict stand is preserved on the Missouri Prairie State Park in Barton County, Missouri (formerly one of the leading prairie hay producing areas). Late estival aspect, September.

FRES No. 39 (Prairie Ecosytem). K-66 (Bluestem Prairie, but as a subunit within tallgrass prairie that did not occur at the mapping scale used by Kucher). The SRM rangeland cover type designated Cordgrass (SRM 726) listed nine variants, but these were all for the Gulf Coast prairies and marshes. S. pectinata was not included in SRM 726. It might seem that the prairie cordgrass community would logically be included as part of the Bluestem Prairie designation (SRM 710), but this species, which forms an obviously distinct climax unit (a consociation), was quite properly not listed thereunder (it is it's own unique rangeland cover type). The SRM (Shiflet, 1994) just misssed it, plain and simple. This is somewhat surprising given the profound impact of Weaver and the University of Nebraska network on generations of rangemen. There should be a Prairie Cordgrass rangeland cover type with a specific and single SRM number and description. Given that the Society (Shiflet, 1994) designated Cordgrass and Sea Oats cover types within the Gulf Coast tallgrass prairie, a Prairie Cordgrass designation and description should be added (if for no other reason than consistenty of presentation). As with the current collection of slides, description of range cover types is an unfinished project. Central Irregular Plains- Cherokee Plains Ecoregion, 40d (Chapman et al., 2002).

96. Prairie cordgrass community- This was an example of the Spartina pectinata consociation that once made up thousands of acres of virgin wet or mesic prairie at the edge of marshes in the Central Lowlands region. It was probably the most productive of all the actual prairie communities with the possible exception of some parts of the bottomland switchgrass-Canada wildrye-eastern gamagrass community (Weaver, 1954, ps. 35-36). Prairie State Park, Barton County, Missouri. Estival aspect, mid-July. FRES No. 39 (Prairie Ecosystem). A subunit of K-66 (Bluestem Prairie).SRM 710 (Bluestem Prairie), or more generally, SRM 601 (Bluestem Prairie). Central Irregular Plains- Cherokee Plains Ecoregion, 40d (Chapman et al., 2002).

97. Inforescence of prairie cordgrass- Even the flower cluster of this productive prairie grass is big, rank, and tough. Weaver (1954, p. 33) explained that both Indians and white pioneers used this species as thatch for lodges, cabins, and even corn cribs. Prairie State Park, Barton County, Missouri. Estival aspect, mid-July.

98. Prairie cordgrass- Flowering stalks of prairie cordgrass (famed Chimney Rock, before latest loss of spire height, in background).

Organization note: further treatment of prairie cordgrass was given below under the section of tallgrass prairie grasses.

Morrill County, Nebraska. July.

99. Tallgrass prairie in its full glory (peak standing crop; maximum morphological development of tallgrasses at anthesis and fruit set)- This scene shows where "tallgrass" gets it's name. A Loamy Bottomland range site in the Smokey Hills physiographic unit of the Central Lowlands in central Kansas dominated by bottomland switchgrass with big bluestem and Indiangrass as associates. Grasses exceed 8 feet in height on this fertile, high water-holding capacity soil. Drier range sites in this area are mixed prairie so tallgrass prairie is postclimax on the alluvial lowland site. The gallery forest growing along a creek in the background is dominated by eastern cottonwood (Populus deltoides) and sycamore (Platanus occidentalis) with bur oak (Quercus macrocarpa) as the major associate.

The Smokey Hills is the geologic-physiogrphic unit immediately west of the Flint Hills which in turn is immediately west of the Osage Questas physiographic unit. Tallgrass prairie is the regional or climatic climax of the latter two and the eastern portion of the Smokely Hills. Geologic ages ago this region was covered by a sea. When the Indians had it to themselves it was a sea of grass.

Tallgrass prairie such as this is, among the herbaceous (not bamboo) grasslands, to the rangeman and grassland ecologist what the redwood forest is to the forester and forest ecologist.This is the ultimate expression of the grassland formation. Lincoln County, Kansas. Late estival aspect, August. FRES No. 39 (Prairie Ecosystem), K-66 (Bluestem Prairie), SRM 710 or 601 both designated as Bluestem Prairie), both the Northern and Southern Great Plains regions in Shiflet (1994) claimed the tallgrass prairies of Kansas (and any fool can see why!). Central Great Plains- Smoky Hills Ecoregion, 27a (Chapman et al., 2001).

100. Autumnal aspect of tallgrass prairie as God and the redman knew it—Upland site dominated by Indiangrass; typical Four Horsemen with willow (Salix sp.), smooth and skunkbush sumac (Rhus glabra, R. trilobata = R. odorata) in draws. Smoky Hills, Maxwell Game Preserve, McPherson County, Kansas. October. FRES No. 39 (Prairie Ecosystem). K-66 (Bluestem Prairie). SRM 710 (Bluestem Prairie) or SRM 601 (Bluestem Prairie). Central Great Plains- Smoky Hills Ecoregion, 27a (Chapman et al., 2001).

101. Another shot of fall aspect of Four Horsemen dominated-tallgrass prairie to show meaning of “tallgrass” relative to a two-year-old Longhorn bull.Wichita Mountains Wildlife Refuge, Commanche County, Oklahoma.October. FRES No. 39 (Prairie Ecosystem). K-66 (Bluestem Prairie). SRM 710 (Bluestem Prairie) variant or SRM 601 (Bluestem Prairie) variant. Central Great Plains- Wichita Mountains Ecoregion, 27k (Woods et al., 2005).

Texas' Grand & Fort Worth Prairie

102. Grand Prairie sampler- Grand Prairie of northcentral Texas in dead of winter. Climax range vegetation presented here was a nearly single-species stand (a consociation if there ever was one) of little bluestem with the major associate species meadow dropseed (Sporobolus asper var. drummondii). Other species, including Indiangrass and Texas wintergrass, were incidential and, basically, non-existent. Range condition class was obviously Excellent.

Green trees were mottes of live oaks (Quercus virginiana var. fusiformis= Q. fusiformis) that had their own unique understorey. This layer(s) of the live oak mottes included cool-season grassses, the dominant species of which were Texas wintergrass and Canada wildrye, warm-season grasses (mostly little bluestem, big bluestem, Indiangrass, and vine mesquite), careic sedges, and shrubs the major one of which was spring herald or elbow-bush (Forestiera pubescens). The Texas live oak motte range type was treated separately in this publication under the heading, Eastern or Southern Live Oak, in Miscellaneous Forest Types under Woodland and Forest Types.

By convention, convenience, and (probably most importantly) practical necessity the various prairies of northcentral Texas havebeen included (lumped in) with the Cross Timbers (see for eg. Diggs et a. 1999, ps. 42-54). Geology and soils of these two natural resource and vegetational units are drastically different with Cross Timbers developing sandy, generally deep soils whereas Grand Prairie soils are underlaid with limestone (often in the form of caliche) and are typically shallower than their sandy land savannah counterparts. In Texas' Cross Timbers and Prairies vegetational (land resource) area, live oak almost always grows only on calcareous (limey) soils so that these evergreen hardwoods are reliable plant indictors of Grand Prairie soils and vegetation.

Across the United States there have been several so-called Grand Prairies including another famous one in Illinois and a small one in Missouri. In Texas in a literal sense (strictly speaking and again by convention) the Grand Prairie as a natural unit of land and vegetation includes the Forth Worth Prairie and Lampasas Cut Plain (Diggs et al., 1999, 48-54). Natural range vegetation of the Grand Prairie, especially of the Forth Worth Prairie portion, is similar to that of the Texas Blackland (=Waxy or Waxyland) Prairie being the "standard" North American tallgrass bluestem prairie but with Texas wintergrass as a cool-season dominant and with considerably more diversity of forbs and woody plants. Kuchler (1964, 1966) maped both the Forth Worth and Blackland Prairies as the same unit, Blackland Prairie (Andropogon-Stipa).

The definitive description of the Fort Worth Prairie portion of the Grand Prairie vegetational unit remains the classic monograph of Dykserthuis (1946).

Erath County, Texas. February, hibernal aspect. FRES No. 39 (Prairie Ecosystem). K-68 (Blackland Prairie), SRM 717 (Little Bluestem-Indiangrass-Texas Wintergrass). Plains Grassland 142.1, "Tall-Grass" Series, specifically Bluestem "Tall-Grass Series, 142.11 of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Cross Timbers- Limestone Cut Plain Ecoregion, 29e (Griffith et al., 2004).

103. Grand Prairie form of tallgrass prairie- The greater Grand Prairie ecosystem, biotic community, etc. historically has been regarded as having the Fort Worth Prairie as one of its ecological units (or subunits) located north of the Brazos River (Dyksterhuis, 1946, fig.1). Most of the Fort Worth Prairie that Dyksterhuis (1946) described has been gobbled up by urban sprawl (including exurban subdivisions). Devout, die-hard conservationists continue to bravely fight a last-stand, battle-to-the-knife siege to save the last remnants of this once magnificant grassland.

Although just to the south of the orginally delineated Fort Worth Prairie (Dyksterhuis, 1946, fig.1), this example of little bluestem-Indiangrass tallgrass prairie presented in these three landscape shots provided students with an example of tallgrass prairie in northcentral Texas that is distinct from tallgrass prairies of Blackland Prairie and Lampasas Cut Plain biotic/ecosystem units. In the hotter climate of this more southern subhumid zone with its generally drier soil profile, Indiangrass replaces big bluestem as the "first-among-equals" dominant on more favorable microsites while little bluestem occupies or performs this role on more typical ("average") environments throughout most of the Southern Tallgrasss Prairie Region. This is distinct from the Northern Tallgrass Prairie Region of Kansas, northern Oklahoma, and Missouri. where big bluestem is king among the Four Horsemen tallgrasses on a zonal (region- wide) scale and on upland mesophytic range sites and where there is more physical (spatial) separation among all four dominant tallgrasses.

Little bluestem was the clear dominant in the first and third of these three photographs with Indiangrass the second most abundant tallgrass and the associate species overall. In the second slide Indiangrass was the dominant tallgrass, but western ragweed had greater foliar cover in some local spots (microsites) as for instance in left foreground. This "species mix" illustrated the frequently found condition in which climax dominants (decreasers) grow immediately adjacent to invaders. This spatial relationship sometimes exist even on virgin range vegetation as on this Excellent condition classs range. There is usually some biologically rational explaination for this seemingly contractionalry ecological state of affairs. The direct explanation in the case of western ragweed beside Indiangrass was that this was a more moist microhabitt (hence Indiangrasss outcompeted the otherwise dominant little bluestem) and buffalo had "played" in the mud creating such a zootic disturbance that Indiangrass was mostly killed out by the pawing, gouging, wallowing, etc. and western ragweed migrated in and achieved dominance on the "new land" created by the disturbance.

The third photograph featured a local patch of an autumnal society of forbs the main species of which were the composites old-field or gray goldenrod (Solidago nemoralis var. longipetiolata), plains blackfoot (Melampodium leucanthum), and narrowleaf gayfeather (Liatris mucronata). This local society was presented in the next photograph.

All trees in these three "photoquadrants" were canyon live oak (Quescus virginiana var fusiformis= Q. fusiformis) which on Grand Prairie are restricted to calcareous (having some form of calcium such as calcium carbonate, like caliche, or lime) soils.

Somervell County, Texas. Mid-October; late estival aspect, peak standing crop just before onset of winter dormancy.FRES No. 39 (Prairie Ecosystem). K-68 (Blackland Prairie), SRM 717 (Little Bluestem-Indiangrass-Texas Wintergrass). Plains Grassland 142.1, Bluestem "Tall-Grass" Series, specifically "Tall-Grass Series, 142.11 of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Cross Timbers- Grand Prairie Ecoregion, 29d (Griffith et al., 2004).

104. Late season gathering- Autumnal society of forbs on a little bluestem-Indiangrass climax range type in the Grand Prairie Area of northcentral Texas. The major forbs were members of the Tubuliflorae subfamily of Compositae: plains blackfoot, old-field or grey goldenrod, and littleleaf gayfeather. This local (locally restricted or small) community was shown in relation to surrounding range vegetation in the immediately preceding photograph.

Society is the term and concept coined by Clements (1916; ps. 130-134). This is an example of an aspect society, one determined basically by seasonality (eg. autumnal aspect thus autumnal society in the example shown here). Later, Clements (1935, p. 276) proposed the term of sociation for a climax aspect society and socies for a seral aspect society. It is easy to see how F.E. Clements "turned off" some ecologists (maybe most who were not Clements' disciples) with his extreme verbage and coinage of terms. Notwithstanding, the idea of plant society is still a useful one--as long as the baggage of terminology does not weight it down.

In the more recent conceptual view of Landscape Ecology the local community (microcommunity) of forbs can be seen as a small patch within the matrix of the surrounding tallgrass (little bluestem-Indiangrass) community (macrocommunity). Clements did not have a monopy on ecological vocabulary.

105. Range vegetation along a tallgrass prairie stream- A small, semi-permanent (dry only in drought) stream of brook-size on the tallgrass prairie range shown in the preceding three-slide set was bais of two ecosystems: 1) wet prairie and 2) woody riparian, both of notable composition and structure. On the riparian and adjacent moist prairie zones taken together a savanna had developed made up of mesic tallgrass species with pecan (Carya illinoinensis), sugarberry (Celtis laevigata), cedar elm (Ulmus crassifolia), and American elm (U. americana)--in that order except that American elm was more abundant deeper in interior of the woody riparian woodland--dominating the woody layers and completing the savanna physiogonomy. In addition to the tree component, the woody plant phase also included a well-developed shrub layer comprised of possum-haw holly (Ilex decidua), elbow-bush (Forestiera pubescens), buttonbush (Cephalanthus occidentalis), common greenbriar (Smilax bona-nox), and chittamwood (Bumelia langulosa var. oblongifolia). Post, rock or blueberry cedar or juniper (Juniperus asheii) had started to invade, an inevitable on-going process in absence of prairie fire.

The wet prairie extended from an herbaceous riparian zone that was devoid of a woody component and extended outward to the little bluestem-Indiangrass tallgrass prairie that was covered immediately above. Species composition of this wet prairie was dominated by bottomland switchgrass with Indiangrass and little bluestem as associates along with the following other grasses (in this approximate order): bushy bluestem (Andropogon glomeratus),. sideoats grama, two dropseeds (S. clandestinus, purpleflower dropseed, primarily; Sporobolus vaginflorus, poverty dropseed, secondarily), Canada wildrye, seep muhly (Muhlenbergia reverchonii), broadleaf woodoats (Uniola latifolia), purpletop (Tridens flavus), King Ranch bluestem (Andropogon ischaemum), and Johnsongrass (Sorghum halapense). This was a pristine stand of wet prairie and the two exotic grasses (latter two species) were not competitive with the native tall- and mid-grasses. Main forbs were Illinois bundleflower, frostweed or white crownbeard (Verbesina virginica), curlycup gumweed (Grindelia squarrosa), and blue boneset (Eupatorium coelestinum). The two important shrubs were buttonbush and indigobush (Amorpha fruticosa). An unidenifiable (no flowers or fruit present) species of spikerush (Eleocharis sp.) was the sole vascular plant actually growing in the stream (bottom of stream channel). Pecan, sugarberry, and cedar elm were the three tree species tht grew in the wet prairie. They were also the only trees that grew beyond the riparian (stream bank) zone.

Most of he riparian zone was more of a woodland plant community with an herbaceous layer of grasses and forbs along with a tree and shrub layer. This riparian vegetation was treated in the slide and caption immediately following this treatment.

Somervell County, Texas. Mid-October; late estival aspect, peak standing crop just before onset of winter dormancy.FRES No. 39 (Prairie Ecosystem). K-68 (Blackland Prairie), Local mesic savanna vriant of SRM 717 (Little Bluestem-Indiangrass-Texas Wintergrass). Wet prairie or riparian variant of "Tall-Grass" Series, specifically Plains Grassland 142.1, Bluestem "Tall-Grass Series, 142.11 of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Cross Timbers- Grand Prairie Ecoregion, 29d (Griffith et al., 2004).

106. Nice place for a prairie picnic- Interior of a bottomland forest or woodland that developed on the riparian zone of a brook-sized, semi-permanent (dry in drought) stream flowing through a little bluestem-Indiangrass tallgrass prairie in the Grand prairie Region of northcentral Texas. This range vegetation had developed downstream from the grassland-savanna vegetation presented in the preceding three-slide set (to immediate left of scenes shown in these three preceding photographs). The large tree (foreground) was an old, fire-scarred pecan. The tree in background was a cedar elm. Major herbaceous species included Canada wildrye, purpletop, little bluestem, broadleaf woodoats, and Texas wintergrass. The most abundant forb was frostweed or white crownbeard. The major shrub was possomhaw holly; other shrubs included elbow-bush, buttonbush, and invading (due to fire-cessation) blueberry, post, or Ashe juniper.

Note on organization and location: a climax, mixed hardwood (sugarberry-cedar elm-pecan) forest with an understorey dominated by Canada wildrye and broadleaf woodoats that developed on the Brazos River floodplain was presented under Forests and Woodlands in the chapter entitled Miscellaneous Forests.

This narrow band of bottomland forest that developed along the stream was a corridor (one of the three major elements in Landscape Ecology) within a matrix (a second major element in the Landscape Ecology conceptual view) of tallgrass prairie. This forest range cover type was included at this point in treatment of tallgrass prairie to show the connectedness of native plant communities, help students grasp the continuity of natural vegetationn, and provide a more complete veiw of range vegetation within the Tallgrass Prairie Region.

Somervell County, Texas. Mid-October; late estival aspect, peak standing crop just before onset of winter dormancy.FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-92 (Elm-Ash Forest). SAF 93 (Sugarberry-American Elm-Green Ash) and/or SAF 94 (Sycamore-Sweetgum-American Elm). Absence of sycamore and sweetgum and presence of sugarberry suggested that SAF 93 was more appropraite; however, the former cover type designation for SAF 94 included pecan so choice was unclear.Southeastern Swamp and Riparian Forest 223.1, Mixed Hardwood Series, 223.13 (Brown et al., 1998, p. 43). Cross Timbers- Grand Prairie Ecoregion, 29d (Griffith et al., 2004).

107. Two on Texas tallgrass prairie- A Grand Prairie tallgrass prairie range dominated by--in fact, a consociation of--Indiangrass and behind it a mixed hardwood bottomland forest of eastern cottonwood (Populus deltoides subsp.deltoides) and cedar elm (Ulmus crassifolia), co-dominants, with common or saw greenbriar (Smilax bona-nox) as the associate that had developed along a small stream.

The associate (actually, more like the most abundant minor) grass species of the tallgrass prairie was purpletop which had greatest cover at edge of grassland and forest. The most common prairie forb was tall goldenrod (Solidago altissima). There were a few stray plants of King Ranch bluestem (Andropogon ischaemum= Bothriochloa ischaemum var. songarica), a dreadful weed on the virgin sod of this climax vegetation.

Other abundant species of the cottonwood-cedar elm-greenbriar forest included trumpet creeper (Campsis radicans), poison oak/poison ivy (Rhus radicans= R. toxicodendron= Toxicodendron radicans), and giant ragweed (Ambrosia trifida). There were some small trees of sugarberry (Celtis leavagada), but these were not enough to include in descriptive name of the existing forest. Work by Rosiere et al. (2013) showed that sugarberry is the first dominant tree species in some mixed hardwood forests that develop along larger streams in northcentral Texas. Such was not the situation here; nor was pecan (Carya illinoinensis) present in sny detectable portion along the gallery forest of this prairie stream.

Hamilton County, Texas. Mid-October; late estival aspect, peak standing crop immediate post-anthesis in indiangrass.FRES No. 39 (Prairie Ecosystem). K-68 (Blackland Prairie), Local mesic savanna vriant of SRM 717 (Little Bluestem-Indiangrass-Texas Wintergrass). Wet prairie or riparian variant of Plains Grassland 142.1, "Tall-Grass" Series, specifically Bluestem "Tall-Grass Series, 142.11 of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Cross Timbers- Grand Prairie Ecoregion, 29d (Griffith et al., 2004).

108. So little of what was so much- Relict virgin tallgrass vegetation of the Fort Worth Prairie. The Fort Worth Prairie is (sadly, was) a tallgrass prairie quite similar to the climax tallgrass prairie communities of the Flint Hills, Osage Cuestas, Cherokee Prairie, Northcentral Prairie, and related range vegetation found from northcentral Kansas southward to northcentral Texas. The Fort Worth Prairie at time of the white man's buffalo hide and, later, cattle frontiers extended for roughly 110 miles south of the Red River and was situated between the Eastern Cross Timbers and Western Cross Timbers (Dyksterhuis, 1946). The floristically and vegetationally related Texas Blackland Prairie had developed just east of the Eastern Cross Timbers and was distinct from the Forth Worth and Northcentral Prairies of northern Texas.

The climax range vegetation presented seen here was being grazed by cattle in a grazing management practice that was some variant form of Short Duration Grazing (or, probably more correctly, a hybrid between SDG and a High Intensity Low Frequency scheme). Domestic sheep also grazed this range, but in a free-ranging, continuous grazing practice. The range plant community presented here and in the next four slide-caption sets was in the death grip of a Severe Drought (Palmer Scale). Presence of forbs of pioneering or other low seral stages, especially the weedy composite, annual broomweed (Gutierrezia draculoides) growing beside decreaser tallgrass species (namely little bluestem, Indiangrass, and big bluestemi) spoke ecological volumes of the impact of Severe Drought.

In addition to the three dominant tallgrass species listed above other, major (associate) grasses included Texas cupgrass (Eriochloa sericea), sideoats grama, and Texas wintrgrass (Stipa leucotricha). Hairy grama (Bouteloua hirsuta), silver bluestem (Andropogon saccharoides), and tall dropseed (Sporobolus asper) were other members of the climax vegetation of this bluestem-dominated prairie. Tall dropseed was a minor decreaser while silver bluestem and hairy grama were increaser species. Texas wintergrass, a local climax dominant to associate, was the only major (generally a decreaser) grass that was a cool-season species. Canada wildrye (anothr decreaser) was a minor grass species that was also a cool-season species. The dominant forb was the rhizomatous, pereennial composite western ragweed (Ambrosia psilostachya)-- other than local populations of annual broomweed. Both of these are generally regarded as invaders in this tallgrass prairie climax vegetation.

Parts of this range were being invaded by honey mesquite (Prosopis glandulosa) and sugarberry (Celtis leavigata). The current managers of this and other ranges of this historic ranch were "scared to death of fire", and what the ranges of this ranch needed more than anything was prescribed burning. Texans need to take a lesson on managing tallgrass prairie from Jayhawkers (Kansans), the master prairie burners. Tallgrass prairie simply cannot persist--at least not properly--with all it's functions in the absence of periodic surface fire.

McFarland Ranch, Parker County, Texas. Mid-October; early autumnal aspect at peak standing crop. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 710 (Bluestem Prairie) and/or SRM 717 (Little Bluestem-Indiangrass-Texas Wintergrass): overlap between these two cover types (Shiflet, 1994, ps. 90-91, 95-96). Cold Temperate Grassland 142, Plains Grassland 142.1, Bluestem "Tall-Grass" Series 142.11 (Brown et al, 1998, p. 40). Range site: Shallow (Soil Conservation Service, 1977). Cross Timbers- Grand Prairie ecoregion 29d (Grifith et al., 2004).

109. What a Texas tallgrass prairie should be- Two close-in "photoquadrants" of a relict of the Fort Worth Prairie, a tallgrass prairie floristically and vegetationally related to tallgrass prairies to the north and east, including those of the Flint Hills, Osage Cuestas, and Smokey Hills in Kansas, Oklahoma, and Missouri.

Dominant grasses in these two "photoplots" were (in this order of cover dominance) little bluestem, Indiangrass, and big bluestem. These are three of the four dominant, decreaser, panicoid grasses of the once-immense tallgrass association; switchgrass was the one missing member of the famed Four Horsemen of the Prairies. Switchgrass was rare on this range on which sideoats grama, Texas wintergrass, Texas cupgrass, tall dropseed, and hairy grama were associate grass species depending on local microhabitat.

The major forbs on this Excellent range condition class prairie were western ragweed and, in current conditions of a Severe Drought (Palmer Scale), the pioneer composite, annual broomweed. Other forbs included the annual euphorb, snow-on-the mountain (Euphorbia marginata), and the showy (and rather unique) perennial known as flax-leaf stenosiphon or false tall guara (Stenosiphon linifolius).

There was "early onset" of brush invasion by such woody species as honey mesquite and sugarberry. Current managers of this historic northcentral Texas ranch were afraid to use prescribed burning, and grassland (especially tallgrass prairie in humid and subhumid precipittion zones) cannot be maintained (preserved) without fire. Fire is a natural part of tallgrass prairie habitat and elimination or, even, reduction of this critical component will eventually result in loss of this priceless rangeland cover type. Fire and grassland (again, especially tallgrass prairie) are inseperable. The tallgrss prairie ecosystem evolved under fire.

110. Some of what precious little was left- Two "photoquadrants" of a relict livestock of the once vast Fort Worth Prairie. The ugly sprawl of the Fort Worth-Dallas "metromess" has destroyed much of this once-fine grassland that was described by (Dyksterhuis (1946). The Fort Worth Prairie had parts of the famed cattle market trails or drove roads, the Chisolm Trail and Shawnee Trail. Now much, if not most, of the remnants of these paths of livestock commerce are paved over.

The examples presented here were on the old McFarland Ranch. These two "photoplots" showed big bluestem, Indiangrass, and little bluestem growing side-by-side at peak standing crop. Another decreaser grass species, though with much less cover was Texas cupgrass (Eriochloa serecia). Accompanying forbs included snow-on-the mountain and annual broomweed (Gutierrezia draculoides). These annuals were likely more abundant in one of the later years of a Severe Drought (Palmer Scale). The perennial forb, western ragweed (Ambrosia psilostachya) was also common. This was a small swale habitat such that soil moisture conditions were more favorable than that of contiguous environments. Another forb with considerably less total foliar cover was the giant, perennial member of the snapdragon family (Onagraaceae) known as flax-leaf stenosiphon or false gaura.

Yes indeed, this much biomass and this extent of grass growth had taken place during what else but another drought and with beef cattle and sheep grazing! It is called Range Management becuse--in final analysis--it is management of grazing resources and not weather that determines condition of the range.

This range was being managed by a form of High Intensity-Low Frequency (or a hybrid of that and Short Duration Grazing) for cattle grazing. As with white-tailed deer, sheep were not contrained by electric fencing used for various cattle-grazing units so these smaller ruminants grazed as free-ranging animals within barbed-wire perimeter (property line) fences.

Management of this range was far from perfect. Current management planners and personnel refused to use prescribed burning which resulted in ongoing nvasion by a number of woody species. Presence of such brush (honey mesquite, ) attested to this neglectful omission of a critical range tool and a pivotal natural factor in the life of tallgrass prairie. Even in the absence of burning, one of Nature's blessings of the prairie, vegetation of this range was in high Good to low Excellent condition class.

McFarland Ranch, Parker County, Texas. Mid- October; early autumnal aspect at peak standing crop. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 710 (Bluestem Prairie) and/or SRM 717 (Little Bluestem-Indiangrass-Texas Wintergrass): overlap between these two cover types (Shiflet, 1994, ps. 90-91, 95-96). Cold Temperate Grassland 142, Plains Grassland 142.1, Bluestem "Tall-Grass" Series 142.11 (Brown et al, 1998, p. 40). Range site: Shallow (Soil Conservation Service, 1977). Cross Timbers- Grand Prairie ecoregion 29d (Grifith et al., 2004).

111. Inside or out: any difference?- Two views of the vegetation of tallgrass prairie cattle and sheep range on the Fort Worth Prairie outside and inside one livestock exclosure. In the first view the range vegetation protected from livestock grazing (vegetation inside the exclosure) was on the left whereas in the second view the range vegetation protected from livestock grazing (vegetation inside the exclosure) was on the right. In the first view there appeared to be greater cover of western ragweed inside the exclosure while in the second view there was greater apparent cover of tallgrass species (little bluestem, Indiangrass, and big bluestem in tht relative order) inside the exclosure.

Such apparent (visual or estimated; not measured) differences in cover and/or biomass were very restricted within the exclosure. The only obvious or undeniable difference was the near absence (essentially zero cover of annual broomweed inside the livestock exclosure. White-tailed deer also foraged on this range and easily could have jumped the three and a half foot tall wovern wire fence.

Sheep were permitted to move freely from one pasture to another on this ranch, but beef cattle were managed by a High Intensity Low Frequency (or a hybrid of this HILF scheme and Short Duration Grazing) grazing practice.

The range vegetation seen here at peak standing crop (point of maximum yield or biomass production) had grown during the second year of Severe Drought. Range can survive and produce a good grazing crop under all but the most extreme or harsh of growing conditions. Give it a chance.

McFarland Ranch, Parker County, Texas. Mid- October; early autumnal aspect at peak standing crop. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 710 (Bluestem Prairie) and/or SRM 717 (Little Bluestem-Indiangrass-Texas Wintergrass): overlap between these two cover types (Shiflet, 1994, ps. 90-91, 95-96). Cold Temperate Grassland 142, Plains Grassland 142.1, Bluestem "Tall-Grass" Series 142.11 (Brown et al, 1998, p. 40). Range site: Shallow (Soil Conservation Service, 1977). Cross Timbers- Grand Prairie ecoregion 29d (Grifith et al., 2004).

112. God's own tallgrass bouquet- Three views at progreessively closer camera distance of the inflorescences of the three dominant tallgrass species on a cattle-sheep range on the Fort Worth Prairie in northcentral Texas at peak standing crop and full-bloom stage. One plant each of little bluestem, Indiangrass, and big bluestem (the three major species) were growing in such close proximity to each other that the photographer could bend sexual shoots of each plant (species) together or one shoot atop the others for these photographs.

When these photographs had been taken the photographer released the shoots unharmed to their original (pre-photographs) position. This showed the close distance of these plants (species) to each other and gave students an idea of spatial relations of them. Little bluestem was the dominant of this climax tallgrass community while Indiangrass and big bluestem were local associate species. The overall associate species in this tallgrass prairie climax community was sideoats grama, but there were places (microsites) were Indiangrass or, less commonly, big bluestem were associate species to little bluestem.

These plants were on a cattle-sheep range where they were subject to grazing by these two domestic species as well as by white-tailed deer. The shoots shown here had grown during a Severe Drought (second year of Severe Drought in a row).

McFarland Ranch, Parker County, Texas. Mid- October; early autumnal aspect at peak standing crop. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 710 (Bluestem Prairie) and/or SRM 717 (Little Bluestem-Indiangrass-Texas Wintergrass): overlap between these two cover types (Shiflet, 1994, ps. 90-91, 95-96). Cold Temperate Grassland 142, Plains Grassland 142.1, Bluestem "Tall-Grass" Series 142.11 (Brown et al, 1998, p. 40). Range site: Shallow (Soil Conservation Service, 1977). Cross Timbers- Grand Prairie ecoregion 29d (Grifith et al., 2004).

Texas' Northcentral (Red River) Prairie

The following section was an example of tallgrass prairie in northcentral Texas. Similarly to other prairies, such as Texas' Grnd Prairie, this area of grassland has at least something of a name being designated as the Northcentral Prairie. The grassland was a form of transition from the Central Lowlands to the Rolling Red Prairie adjoining it to the west. It was in the greater floodplain of the Red River being drained by the Little Wichita River into Red River. Specifically, this land area is part of the Osage section of the Central Lowlands physiographic province south of the Arbuckle-Wichita Montains uplift (Fenneman, 1938, ps. 609-610, 619-620). The Wichita is the principal geologic formation (Gordon, 1913, ps. 22-27, 44-45).

The portion of the Northcentral Prairie treated below had been part of a historic ranch owned and managed by the J.L. Hudgins and Bryant Edwards families for well over a century. Some of the grassland of the original Edwards Ranch along with the former Earl Springs Ranch was owned and being managed as the F Bar Springs Ranch (over 12, 000 acres) by Diaz and Laura Murray for 14 years up to the time of these slides.

Other parts of the Edwards outfit had been owned and operated by Emry Birdwell and Deborah Clark for 11 years prior to time of photographs. The famed--and well managed--ranch consisted of 14,200 acres. The Birdwell & Clark Ranch website described the grazing management program as originally consisting of three grazing or pasture cells (each of about 4,000 acres and stocked with 1500 to 1800 head of stocker cattle) for a total of 140 paddocks (varying from 45 to 145 acres). These paddocks had been developed around and extended outward from water sources Three herds of 1500-1800 head each grazed through various paddocks in each cell.

During the drought of 2011 to 2014 the Birdwell & Clark Ranch modified its original grazing method and changed from a three-cell/three herd schedule to a one-cell/one herd (of up to 3,300 head) scheme. This grazing method was a form of Short Duration Grazing. It was the grazing mangement arrangement at time of the following photographs which were taken one year after the 2011-2014 drought. Photographs were of tallgrass prairie vegetation at peak standing crop and end of the warm-growing season following removal of stockers, a rest or recovery period which had permitted (encouraged?) regrowth of grass plants to size, vigor, and yield similar to that of an ungrazed prairie.

This was one of the finest cases of sound range stewartship on tallgrass prairie ever viewed by your author. It was uplifting to be able to present to students this sterling esample of sound range management, efficient livestock production and profitable ranch operation on one of the most productive grassland range types in North America.

113. Try to top this- Tallgrass prairie range in the Northcentral Prairie just below Red River managed as a padddock in a short duration grazing arrangement. Local co-dominants in these two views were switchgrass and Indiangrass. Overall the lead dominant was little bluestem. Other important grass species included the naturalized King Ranch bluestem (Andropogon ischaemum= Bothriochloa ischaemum) silver bluestem, sideoats grama, thinseed paspalum (Paspalum setaceum), meadow dropseed (Sporobolus asper var. drummondii), white tridens (Tridens albescens), hairy grama, blue grama, and buffalograss. Big bluestem was a minor component, and it was this lesser cover of big bluestem that distinguished this subhumid southern portion of the tallgrss prairie region from tallgrass prairie vegetation ranging from central Oklahoma northward to the Greak Lakes prairies.

The diversity of grass species of this tallgrass prairie stocker range was tremendous in going from tallgrass through midgrass down to shhortgrass species. This is a species "lineup" rivaling that of mixed prairie grassland of the adjoining Rolling Red Plains to the west. This was undeniably tallgrass prairie, however, due to the limited cover, density, etc. of the midgrass and shortgrass species. Even relative cover of the generally plentiful sideoats grama and silver bluestem, co-dominant species of the Rolling Redlands mixed prairie, was of obviously secondary importance to tallgrass species.

The prinncipal forb was "far and away" western ragweed (Ambrosia psilostachya). Most of the widely sdattered forbs were composites like narraowleaf gayfeather (Liatris mucronata) and elegant gayfeather (L. elegans) and legumes such as white prairie clover (Petalostemon candidus).

Brian Edwards (Birdwell & Clark) Ranch, Clay County, Texas. Early October; early autumnal aspect, peak standing crop. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 710 (Bluestem Prairie). Cold Temperate Grassland 142, Plains Grassland 142.1, Bluestem "Tall-Grass" Series 142.11 (Brown et al, 1998, p. 40). Range site: Loamy Prairie (Soil Conservation Service, 1980d). Central Great Plains- Limestone Plains ecoregion 27j (Grifith et al., 2004).

114. "If Quanah Parker came back, I hope he would say this looks like it did before." (Emry Birdwell)- It would appear that is indeed what the great half-breed chief of the Comanche would say. Tallgrass prairie used as range for stocker cattle managedusing a short duration grazing method. This was a closer view of the sward of the pasture introduced in the preceding two-slide/caption unit. Switchgrass (an upland ecotype) and Indiangrass were co-dominants. Little bluestem was the local associate species, but overall it was the most abundant grass. Other grass species included King Ranch bluestem, silver bluestem, sideoats grama, thinseed paspalum, meadow dropseed, big bluestem, white tridens, hairy grama, blue grama, and buffalograss. Based on presence of tall-, mid-, and shortgrass species this grassland had a mixed prairie makeup, but the mid- and shortgrasses were "minor players" on what was conspicuously a tallgrass prairie communiy. Forbs were sparse on this tallgrass prairie.

It was also obvious that this was the climax grassland vegetation so that the range was in Excellent range condition class.

Yes, I believe Quanah Parker would approve of the current management of this former Comanche buffalo range. People's love of the land shines through whether redman, white, or otherwise.

Brian Edwards (Birdwell & Clark) Ranch, Clay County, Texas. Early October; early autumnal aspect, peak standing crop. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 710 (Bluestem Prairie). Cold Temperate Grassland 142, Plains Grassland 142.1, Bluestem "Tall-Grass" Series 142.11 (Brown et al, 1998, p. 40). Range site: Loamy Prairie (Soil Conservation Service, 1980d). Central Great Plains-Limestone Plains ecoregion 27j (Grifith et al., 2004).

115. Through drought, fire and cattle- Tallgrass prairie range four years after the "double whammy" of prolonged drought and prairie fire wildfire on top of grazing by cow/calf pairs in the Northcentral Prairie of north Texas. Four years prior to time of these photographs a wild fire in mid-August burnt this already drought-stressed prairie. There had relatively little recovery of the vegetation the first three years--on-going drought years--post fire. At end of the fourth warm-growing season after the hot summer fire the prairie grasses were staging a comeback.

Apparently the summer fire did not kill existing grass plants (at least not most of them), but stressed them so as to reduce their productivity (of course only herbage or aboveground biomass could be observed directly). After four growing seasons, grass recovery had progressed to herbage yields and returned to the rnage plant community seen here. In other words, while seral species ("weedy" forbs for instance) did increase relative to grasses following the summer fire, this disturbance--and, again, accompanied by Severe to Extreme Drought--was not so harsh as to create classic "old field" conditions so that the climax grassland had to undergo secondary plant succession. Certainly there was some degree of range retrogression, but most revegetation was through recovery of pre-fire plants. Plants of Indiangrasss and little bluestem as seen in the second slide did not start from seed in the current year.

In addition to the potential natural (climax) co-dominants little bluestem and Indiangrass other grasses present included King Ranch bluestem, silver bluestem, sideoats grama, purple threeawn, white tridens, meadow dropseed, purple-flower dropseed (Sporobolus clandestinus), hairy grama, plains lovegrass (Eragrostis intrmedia), and buffalograss. Forbs included such composites as western ragweed, elegant gayfeather, and annual broomweed (Gutierrezia draculoides).

Brian Edwards Ranch (now, Murray F Bar Springs Ranch ), Clay County, Texas. Early October; early autumnal aspect, peak standing crop. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 710 (Bluestem Prairie). Cold Temperate Grassland 142, Plains Grassland 142.1, Bluestem "Tall-Grass" Series 142.11 (Brown et al, 1998, p. 40). Range site: Claypan Prairie (Soil Conservation Service, 1980d). Central Great Plains-Limestone Plains ecoregion 27j (Grifith et al., 2004).

116. Back after typical tallgrass trials- Fours years after a summer wildfire and prolongued drought and proper grazing by cow/calf pairs this tallgrass prairie had recovered to a great degree. There was very limited revegetation of this grassland until the current fourth year following fire. Lingering drought had been a (probably the) major factor that limited grass recovery. Recovery was apparently mostly restoration of vigor and productivity of grass plants living at time of the summer fire.

Grass species seen here included little bluestem, Indiangrass, purpleflower dropseed, King Ranch bluestem, meadow dropseed, sideoats grama, hairy grama, silver bluestem, purple threeawn, plains lovegrass, and buffalograss. Although this was clearly tallgrass and not midgrass or mixed prairie, big bluestem was not encountered. In this subhumid (approximately 30 to 32 inch average precipitation belt) zone big bluestem is not abundant. This is in contrast to tallgrass prairies farther to the north and east in the tallgrass belt of the Central Lowland physiographic province (in which this grassland clearly falls) where big bluestem is the overall regional dominant.

The paucity of range forbs--by criteria of both species and cover--was also noteworthy. Western ragweed was the major forb, but cover and density of this major composite were very limited. This had to be due, to some degree, to the excellent grazing management of this and the other ranges on this well-run, profitable ranch.

This outfit specialized in production what has been known variously as "grassfat beef", "grassfed beef", "grassfat cattle" etc. F Bar Springs Ranch is marketed through or in conjuction with the Grassfed Livestock Alliance of Texas.

Grassfat cattle was, of course, the original or first way (production system) growing meat, especially on range. Cattle (or sheep) were often fattened or finished on their way (as they were slowly driven) to markets. In trail drive days such cattle were often old (as in four or five up to ten or more years old). A human generation or two ago home-raised beef for consumption on family ranches--the family "butcher steer"-- was upwards of three (or more) years old. The beef from such cattle might or might not be tender (cattle were rarely of A maturity). This home-grown, grassfat beef definitely had a "grassy" or "gamey" taste and was characterized by yellow (high Vitamin A content) fat. It had (has) a distinctive flavor which some people liked, got used to, or did not anything different.

Anyway, it was essential to have "good feed" for production of grass- (versus grain-) finished beef. The ranges (most were in Excellent range condition class) on this specialty ranch reflected the basic requirement for "good feed growing on the range". High-quality products (beef, in this instance) from the range cannot be produced on low-quality range.

117. Fall foliage of a tallgrass prairie- The strikingly colorful herbage of a more open portion of a post oak (Quercus stellata)-bluestem/ Indiangrass savannah form of tallgrass prairie on the Nothcentral Prairie just south (yet elevationally above) Red River. This was near the westernmost edge of tallgrass prairie just before moving westward into the mixed prairie of the Rolling Red Plains. Little bluestem was the dominant species with Indiangrass a close associate and, sometimes, co-dominant species. Sideoats grama, hairy grama, silver bluestem, purple-flower dropseed, purple threeawn, and white tridens were other, though generally minor, grasses. Forbs were represented by the rhizomatous perennial, western ragweed, the forb species with greatest cover, and by the oppurtinistic annual, camphorweed (Heterotheca submaxillaris).

This was a cow-calf range plus an abundance of white-tailed deer along with lesser fauna including a array of insects.

This was obviously the climax range vegetation which is regarded by some prairiemen as being most beautiful when in fall coloration (other prairie fans opt for the spring and summer bluegreen of bluestem). Eitherway, it's always beautiful to grasslanders. New England hardwood forests are gorgeous in their famed fall colors, but to prairiemen New England coloration "ain't got nothin'" on an autumn tallgrass prairie.

Brian Edwards Ranch (now, Murray F Bar Springs spring and summer Ranch ), Clay County, Texas. Early October; early autumnal aspect, peak standing crop. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 710 (Bluestem Prairie). Cold Temperate Grassland 142, Plains Grassland 142.1, Bluestem "Tall-Grass" Series 142.11 (Brown et al, 1998, p. 40). Range site: Sandy Loam (Soil Conservation Service, 1980d). Central Great Plains-Limestone Plains ecoregion 27j (Grifith et al., 2004).

118. Color contrast of an autumn savannah- The dark green of post oak leaves at end of their lives (end of the annual growing season) contrasted sharply with the golden colors of "ripening" (=senescing or maturing) grass shoots on a savannah form of tallgrass prairie in the Northcentral Prairies. This distinctive area of tallgrass prairie was in the Red River vicinity (north and south of the stateline river) that is part of the southern Central Lowlands physiographic province. Near its western margin where tallgrass country meets mixed prairie at the eastern edge of the Great Plains post oaks are scrubby compared to those of the Post Oak Savanna and Cross Timbers vegetational areas.

The grasses, by contrast, still reach their tallgrass potential. This was shown by the robust specimen of Indiangrass presented in center foreground in the first slide and lower left corner of the second slide and by little bluestem seen as numerous bunches or tufts in the second slide. (The next--immediately following--set of two slides also presented the typical growth size of these two dominant tallgrass species). Other grasses in these two "photo-plots" included sideoats grama, silver bluestem, purple-flower dropseed, purple threeawn, and plains lovegrass. A common forb was the opportunistic annual composite, camphorweed.

Brian Edwards Ranch (now, Murray F Bar Springs Ranch ), Clay County, Texas. Early October; early autumnal aspect, peak standing crop. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 710 (Bluestem Prairie). Cold Temperate Grassland 142, Plains Grassland 142.1, Bluestem "Tall-Grass" Series 142.11 (Brown et al, 1998, p. 40). Range site: Sandy Loam (Soil Conservation Service, 1980d). Central Great Plains-Limestone Plains ecoregion 27j (Grifith et al., 2004).

119. Beats New England fall foliage (at least to a prairieman)- To rangemen bonded to tallgrass prairie the blazoned fall colors of the prairie plants rivals anything trees of the New England forests put forth. Throw into this climax range vegetation the scattered, scrubby post oaks with an occasional forb like the opportunistic species known as eastern daisy fleabane (Erigeron annuum) with its numerous white heads (right-center foreground of first slide) and mid-continent prairies have a distinctive beauty all their own.

The beauty of such grassland communities was shown at closer camera distance in the second slide which featured little bluestem and Indiangrass, the co-dominants of a Northcentral Prairie range in Excellent range condition class. This prairie pasture was one of those used for production of high-quality grassfed beef that has been skillfully marketed by the Murray F Bar Springs Ranch. Production of a specialty product like grassfat beef requires high-quality pasture like that presented in these images.

Of course, the late-maturity, going-dormant herbage (senescing or end-of-life shoots) seen here do not make for feed of high nutritive value, but with protein supplements this range feed is more than adequate provender for dry pregnant beef cows. This was an ideal example of the necessary end-of-season stage for the tallgrass crop. (Actually, degree of use could have been substantially greater than that of the almost unused appearance of this herbage, but degree of use at one given point of time is not the only or, even, the best measure of utilization. Furthermore, range plants on this ranch had just been through a multi-year drought (and some ranges on this outfit had undergone an August wildfire four years earlier combined with prolonged droguht) such that a little extra herbage growth in the first decent year in quite a spell was the same as money in the bank, a little "velvet" on the range as it were.

Brian Edwards Ranch (now, Murray F Bar Springs Ranch ), Clay County, Texas. Early October; early autumnal aspect, peak standing crop. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 710 (Bluestem Prairie). Cold Temperate Grassland 142, Plains Grassland 142.1, Bluestem "Tall-Grass" Series 142.11 (Brown et al, 1998, p. 40). Range site: Sandy Loam (Soil Conservation Service, 1980d). Central Great Plains-Limestone Plains ecoregion 27j (Grifith et al., 2004).

Subhumid Rolling Redlands or Red Plains Prairie

The set of slides presented in this section were of a tallgrass prairie in a portion of the Great Plains known variously as the Rolling Red Plains, Rolling Redlands, and Reddish Prairie. More specifically from a physiographic standpint this sample of tallgrass prairie was in the Red Hills portion of the Plains Border or Break of the Plains section of the Great Plains physiographic province (Fenneman, 1931, ps. 25-30; Fenneman, 1938, p. 606) or, perhaps, more specifically the Gypsum Hills portion of the Osage section of the Central Lowlands physiographic province that is the southern end of the Red Hills portion of the Great Plains (Fenneman, 1938, ps. 618-619).

This area is on the western edge of the subhumid zone with average annual precipitation ranging from an average of 24 to 30 inches (Oklahoma Climatological Survey, online). This tallgrass prairie was at the westernmost margin of the tallgrass range type. It could be interpreted as the subclimax Andropogon associes (association is a unit of climax; associes, of subclimax) as viewed by Clements (1920, p. 131-134) or as subclimax in development and as postclimax to the true prairie as explained in Weaver and Clements (1938, ps. 520-521). Clements (1920, p. 132) explained that the 30 inch isohyete ("precipitation line") was "[t]he western limit of the subclimax prairie".

It was explained in photo-captions below that tallgrasss, midgrass, and shortgrass species were present in this grassland range, but midgrasses (mostly sideoats grama and silver bluestem) were minor associate species while shortgrasses were "just there" as incidental species. At cursory visual scale ranging from "drive-by" views (physiogonomic appearances) to walk-through and "down on hands and knees" close looks this pristine grassland was tallgrass and not mixed prairie. It was tallgrass prairie in an area so near to the 100th meridian that midgrass prairie would, perhaps, be typically expected.

It was notable that tallgrass prairie extended this far to the west where it met mixed prairie. This example was about midway between the 99th and the 100th meridians. Isohyets (map lines of connecting points having the same, or very similar, amonunts of average annual precipitation) revealed the explanation: this example grassland was in the subhumid not the semiarid zone.

These photographic samples were taken on the Black Kettle National Grassland in Roger Mills County, Oklahoma.

120. Grand sweep of Rolling Redlands tallgrasss prairie- Two landscape scale views of a tallgrass prairie that had developed in the Red Hills-Gypsum Hills of the Great Plains-Central Lowlands transition. Grassland vegetation presented in these sweeping views was in a more mesic area of the subhumid zone at the Prairie-Plains Border of the Break of the Plains. West of where "it ought to be" (ie. a westernmost extension of tallgrass prairie into the Mixed Prairie Region), this grassland appeared as an outlier of tallgrass prairie more typical of the Smoky Hills-Flint Hills-Osage Plains complex.

Indiangrass, which is less mesic or more xeric than big bluestem and switchgrass, was the overall dominant of the tallgrass prairie presented here and in the next four slide/caption sets. Switchgrass, the most mesic of the Four Horsemen of the Prairies (big bluestem, Indiangrass, and little bluestem being the other three), was sparse in these two views of this subhumid form of tallgrass prairie. In addition to the major three tallgrasses, the midgrasses silver bluestem and sideoats grama (overall co-domiants of much of the mixed prairie to the west) were important species. Tall dropseed (Sporobolus asper complex) was locally important and, as shown below, prairie cordgrass was the dominant in low-lying, wet habitats. Shortgrass species hairy grama (Bouteloua hirsuta), blue grama (B. gracilis), and bufffalograss (Buchloe dactyloides) were minor components of this tallgrass prairie.

Major forbs were the legumes, Illinois bundleflower, white and purple prairie clover, and broadtop or bigtop dalea or nine-anther prairie clover (Dalea enneandra). Among the composites, Engelmann daisy (Engelmannia pinnatifida) was most important yet it was "nowheres" a major species. It was noteworthy that Engelmann daisy was at peak bloom on this grassland in early July. By comparison blooming time of Engelmann daisy was two to two and a half months later at this loction than in tallgrass prairies in northcentral Texas where Engelmann daisy is a major spring-blooming forb.

The most abundant (important) shrub was smooth sumac (Rhus glabra), a highly rhizomatous species that grew in small colonies. Plains yucca or soapweed yucca (Yucca glauca) was also frequent though not abundant. (Y. glauca can be viewed as either forb or shrub depending on perception or interpretation.) Another well-dsitributed shrub in this tallgrass grassland was catclaw mimosa or wait-a-minute bush (Mimosa biuncifera).

Black Kettle National Grassland, Roger Mills County, Oklahoma. Early July; early estival aspect. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 710 (Bluestem Prairie). Cold Temperate Grassland 142, Plains Grassland 142.1, Bluestem "Tall-Grass" Series 142.11 (Brown et al, 1998, p. 40). Range site: Loamy Prairie (Soil Conservation Service, 1963, p, 35-36) or Loamy Upland ecological site (Natural Resources Conservation Service, online). Central Great Plains- Rolling Red Hills Ecoregion 27q (Woods et al., 2005).

121. Tallgrasses dominating the Rolling Redlands- Landscape-scale view of tallgrass prairie in the eastern (subhumid) portion of the Rolling Red Plains. In foreground and front part of midground a Loamy Prairie range site was dominated by Indiangrass and, locally, co-dominated with big bluestem.Little bluestem was the major associate species along with Canada wildrye, sideoats grama, and silver bluestem as major grass species. There were scattered plants of hairy grama, blue grama, and bufffalograss but shortgrass cover was a minor component of this grassland. There were plants of Engelmann daisy, Illinois bundleflower, and bigtop prairie clover or broadtop dalea, but the conspicuous forb in foreground was yellow dalea (Dalea aurea).

In the background and back part of midground a Loamy Bottomland range site was dominated by prairie cordgrass and patches of buttonbush (Cephalanthus occidentalis). Switchgrass was present at substantial cover in this mesic Loamy Bottomland range site, which as indicated in the more recent ecological site name of Subirrigated Bottomland, was subirrigated (ie. thr surface aquifer was within rooting depth of plants growing in the soil above this water table).

Black Kettle National Grassland, Roger Mills County, Oklahoma. Early July; early estival aspect. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 710 (Bluestem Prairie). Cold Temperate Grassland 142, Plains Grassland 142.1, Bluestem "Tall-Grass" Series 142.11 (Brown et al, 1998, p. 40). Range site: Loamy Prairie in foreground; Loamy Bottomland in background (Soil Conservation Service, 1963, p, 35-36) or Loamy Upland ecological site Subirrigated Bottomland (Natural Resources Conservation Service, online). Central Great Plains- Rolling Red Hills Ecoregion 27q (Woods et al., 2005).

122. Herbaceous redmen on the Reddish Prairie- Three views of tallgrass prairie in the Rolling Red Plains Region (specifically the Red Hills-Gypsum Hills area) dominated by Indiangrass. Little bluestem was the associate species while big bluestem, prairie cordgrass, silver bluestem, sideoats grama, Canada wildrye, and switchgrass (in roughly that order of foliar cover) were other important grass species in the grassland vegetation shown in these three slides.

There was remarkably little cover of forbs. Illinois bundleflower was the principal forb with catclaw sensitivebrier (Schrankia nuttallii) the first runner-up forb. Both are non-papilionaceous legumes.

123. Representative of tallgrass country at its edge- "Photoquadrant" of tallgrass prairie at its western edge in the Red Hills-Gypsum Hills area of the Great Plains. Dominant range plants were Indiangrass and, locally, smooth sumac. In this "photoquadrant" the associate grass species was Canada or nodding wildrye. Little bluestem was dominant in much of the background, but other major tallgrasses were Indiangrass, big bluestem, and tall dropseed while silver bluestem and sideoats were the principal midgrass species. Forbs were sparse.

124. Happy in the subhumid zone- "Photoplots" of climax range vegetation of tallgrass prairie in the Red Hills-Gypsum Hills portion of the Rolling Red Plains Region. Grassland vegetation in both of these photographic samples was dominated by Indiangrass. The range plant community in the first slide had considerable cover of the leguminous forb, Illinois bundleflower as well as of the leguminous shrub, catclaw mimosa or wait-a-minute bush. Sadly, there was also a small eastern red cedar (Juniperus virginiana) in this first "photoquadrant". Eastern red cdar is a non-sprouting conifer so a good prescribed (or lightening-ignited) fire would handily take out this woody invader (on this range site eastern red cedar is brush, a woody weed or a native ecological invader).

The second "photoquadrant" was of a west-facing slope on which the grassland vegetation was a consociation of Indiangrass. This slope was a road cut that was "hairing over" with the local, climax dominant plant species. Indiangrass was apparently the general area dominant range plant of this westernmost extension of tallgrass prairie. Range site descriptions in the county (Roger Mills, Oklahoma) soil survey (Soil Conservation Service, 1963) listed big bluestem as the potential dominant of most range sites, including Loamy Prairie, the predominant range site featured in this set of slides. Even with his short range reconnaissance this rangeman felt strongly that it was Indiangrass not big bluestem that was the major climax dominant of the less mesic tallgrass prairie. This conclusion was consistent with the rangeland cover type description of tallgrass prairie in northcentral Texas and southcentral Oklahoma where the lower water-requiring little bluestem and Indiangrass and not the more mesic big bluestem define that form of tallgrass prairie (Shiflet, 1994, ps. 95-96). Big bluestem is the general dominant in tallgrass prairies of eastern and central Oklahoma and Kansas, but Indiangrass is the dominant in tallgrass prairies of the western zone of the tallgrass type. The exception is sand bluestem (Andropogon hallii= A. gerardii subsp. hallii) which is the potential dominant of the deeper sands or high-sand range sites of this western portion.

These two views provided a good example of the land surface of the Woodward-Quinlan soil association (Soil Conservation Service, 1963). Parent material for these soils was the Permian Red Beds.

125. On just stick it in- Engelmann daisy (Engelmannia pinnatifida) growing on a tallgrass prairie in the western margin of this type in the Red Hills-Gypsum Hills area of the Great Plains. This was part of the Rolling Red Plains, Rolling Redlands, or Reddish Prairie vegetation zone that extends from northwestern Texas to southwestern Kansas.

General view of shoot (first slide) and capitula inflorescences (second slide). Ray flowers of this member of tribe Heliantheae frequently turn under in this fashion. This can be confusing to observers who are used to seeing these liguliferous flowers extending straight out.

Black Kettle National Grassland, Roger Mills County, Oklahoma. Early July; peak-bloom phenological stage.

126. Tallgrass prairie of Lampasas Cut Plain- Dominance of this plant community by tallgrass species of little bluestem and meadow dropseed qualified this range plant community as an example of interior tallgrass prairie, but strictly speaking this vegetation is part of the Edwards Plateau due to its physiographic affinity with the Great Plains. It was interpreted as part of the Great Plains physiographic province by Fenneman (1931, pgs. 54-59, esp. 55; 1938, ps. 102, 106). Thus this range vegetation was discussed in more detail in the Grassland chapter, entitled Edwards Plateau. This example of prairie was included at this point in interior tallgrass parairie to provide consistency as to physiography, avoid confusion, and provide as much comprehensive coverage within each chapter as was possible.

Lampasas County, Texas. October. Autumnal aspect. FRES No. 39 (Prairie Ecosystem). K-66 (Bluestem Prairie). Variant of SRM 717 (Little Bluestem-Indiangrass-Texas Wintergrass). Cold Temperate Grassland 142, Plains Grassland 142.1, Bluestem "Tall-Grass" Series 142.11 (Brown et al, 1998, p. 40).Cross Timbers- Limestone Cut Plain Ecoregion 29e (Griffith et al., 2004).

127. Sand prairie- An ecotonal tallgrass prairie dominated by little bluestem with yellow Indiangrass and sideoats grama as associate species that developed on deep sand in the West Cross Timbers. This grassland was contiguous with a dwarf forest or shrubland, known locally as sandrough", comprised of sand post oak (Quercus margarettiae) and fiddleleaf greenbrier (Smilx bona-nox). At landscape scale this prairie could be seen as a patch of grassland situated among a matrix of "sandrough".

The conspicuous yellow composite was stiff-leaf goldaster (Heterotheca stenophylla).

Although the soil of this grassland was primarily deep sand it had enough clay and differed (largely in depth and proportion of sand) from the adjoining Patillo-Nimrod-Arenosa soil complex that supported scrub oak trees or large shrubs.

This climax range plant community could fit into either tallgrass prairie (Grand Prairie) or Cross Timbers vegetation so it was included in both chapters, Tallgrass Prairie and Tallgrass Savanna.

Eastland County, Texas. Mid-October; peak standing crop, full-bloom stage to fruit-ripe stage in most plant species.

Loess Hills and Glacial Till Prairies

Perhaps (in all probability) the least common--in fact, rarest--of the tallgrass prairies are those of the best farm land, those whose "highest and best" economic use was field crop production. These include those lands whose soils are loess and glacial till which are some of the richest, most productive soils in North America-- inf not on Earth. The following examples of loess and glacial till prairies were from the few remaining gems of tallgrass prairie ecosystems (although incomplete of the rarer herbivores and all the large carnivores) saved by heroic and dedicated efforts of lone individuals, local groups, state associations up to the deep-pocketed international organizations like The Nature Conservancy. Thank you!

Tallgras prairie in the Western Loess Hills:

About all that remains- Landscape of tallgrass prairie in the loess hills of northwest Iowa. This was a "Four Horsemen of the Prairies" (big bluestem, little bluestem, Indiangrass, and switchgrass) grassland that had never been farmed. Two vegetation and landscape-scale vistas of the largest remaining tract of tallgrass prairi--of virgin sod--in Iowa, a little less than 3000 acres in the Broken Kettle Grassland Preserve "saved" by The Nature Conservancy. In the first of these two slides the conspicuous forb (a legume) in foreground and midground was lead plant (Amorpha canescens) which was growing in a "nearly pure" sward of big bluestem (Andropogon gerardii). Big bluestem is a warm-season, shortshoot tallgrass meaning that its shoots do not elongate until much later in the warm-growing season just before its sexual (flowering) shoots start to elongate. Indiangrass (Sorgastrum nutans) was also preent though at much lower cover and densit;y. Indiangrass is also a warm-season tallgrass, but a longshoot species in that its shoots elongate much earlier in the growing season, long before the flowering sexual shoots elongat. Porcupine grass (Stipa spartea) was another major grass though it is a cool-season species and one often interpreted as more a midgrass than tallgrass species. Porcupinegrass was the major cool-season grass--and another associate to big bluestem, the dominant grass-- in this grassland sward.

The same grass composition--big bluestem, dominant; Indiangrass and porcupinegrass, associates--was in the second (vertical) slidee, and here, too, lead plant was a major player but here, also, in the second"photoplot" compass (Silphium lanceolatum), a perennial composite, was a principal forb challenging its legume counterpart. Most conspicuous in the second slide was local dominance by smooth sumac (Rhus glabra) as obvious in midground. With their well-developed, expansive woody "rootstocks" (rhizomes) Rhus species can spread aggressively. The sumacs are so well adapted to fire that they thrive under frequent--though NOT annual--burning. In saving this priceless gem of tallgrass prairie, for which frequent fire is essential, The Nature Conservancy had wisely applied some prescribed fire. Smooth sumac gratefully responded by prolific sexual sprouting. Such can be a challenge to maintaining the climax tallgrass and midgrass species: adjusting the proper fire regimen for a grassland type that is probably a fire climax (ie. a fire type).

In regards fire, including prescribed burning, on this grassland students should note the presence--undoubtedly an excess cover--of trees. Your reporting rangeman could not help making a comparison of this tree-invaded loess hills tallgrass prairie with the much lower cover of trees (and other woody plants) on tallgrass prairie in the Kansas Flint Hills where annual burning is a ritual neigh on to religious experience.More fire, please.

Broken Kettle Grassland Preserve, Plymouth county, Iowa. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 710 (Bluestem Prairie); SRM 802 (Missouri Prairie.. Plains Grassland 142.1, Bluestem "Tall-Grass" Series, 142.11 of Brown et al. (1998, p. 40). Western Corn Belt Plains 47, Northwest Iowa Loess Prairies 47a Ecoregion (Chapman et al., 2002).

Why it grows like it does- Profile of a loess soil of the Broken kettle Grasslands Preserve. Formation and deposition of loess is a gelogic process to be marveled..

Tallgrass prairie in the glaciaal till plains of the greater Red River Valley:

Tallgrass Prairie in the Lake Agassz Plain- In the repeatedly glaciated northern Central Lowlands physiographic province various tpes of vegetation developed that range from conifer and hardwood forests to aspen parklands to grasslands. These northern grasslands have much the same species ccompositions as grasslands far to the south. The grasslands of the northern glaciated region, including the till plains, include the tallgrass prairie type The example used here was in the Red River Prairie subsection of the Red River Valley section of the Prairie Parkland province as shown in the Minnesota Ecological Land Classification Hierarchy (Minnesota Department of Natural Resources, online). This particular tallgrass prairie was dominated by native warm-season perennial grasses, especially the panicoid grasses. Native cool-season grasses (the festucoid tribes), including Stipa, Poa, and Elymus species, were also present as were egrostoid grasses like Sporobolus and Bouteloua species.

Grasslike species, especially caric sedges (Carex spp.), a wide arrray of forbs, and even a few woody plants combine with the Gramineae (grass family) to produce a range plant community that has an amazing species diversity in spite of a simple, rather nondescript physiogonomy and a simplier vegetaqtional structure compared to neighboring forests.and savannahs.

The tallgrass prairie vegetation in these slides and those below were of the vernal society-- the seasonal plant community made up of spring (cool-season or early warm season; "early blooming) species and later maturing warm-season species in less-mature phenological stages. One of the more prominent forb with it's off-white flowerr cluster was purple meadow-rue (Thalictrum dasycarpum). Other forbs included golden Alexander (Zizia aurea), smooth scouring rush (Equisetum laevigatum) northern bedstraw (Galium boreale), Canada windflower (Anemone canadensis), and Indian hemp (Apocynum cannabium). Therr was also an immature goldenrod (Solidago sp.) that could not be identified. Shrubs were preresented by the woody legume, false wild indigo or false indigo bush (Amorpha fruticosa) and the small shrub, prairie or Arkansas rose (Rosa arkansana), was also present.

Seasonal changes from a physiogonomy as well as a color perspective are more pronounced in the tallgrass prairie--and, perhaps even more so, in the mixed prairie--than in forests. Changes in plant (shoot) height, for example, exhibit more seasonal variation in grasslands, especially tallgrass-dominated grasslands, than those of forest or shrublands. At these same locations, appearances of the range plant community presented in these spring slides (including the ones below) would differ markedly from slides taken in summer and autmmn. Students new to grasslands must bear this in mind when viewing and discussing tallgrass prairie. Eight foot shoots of big bluestem and Indiangrass are not going to appear in photographs of the vernal society of a tallgrass prairie.

The large rocks or small boulders are glacial erratics. The geologic history (if "history" be the word) of this area is fascinating. Students can find numerous referernces on, say, the Ice Ages. Geologists, especially glaciologists, are generally agreed that there have been five major Ice Ages. The most recent (the one we are in now) is the Quaternary or Pleistocene glaciation within which is the current interglacial period, The most recent glacial period was the Wisconsin Glacial Episode or the Wisconsin Glaciation which ended approximately 11,000 to 10,000 years ago. That was probably when the glacial erratics shown here were deposited.

This area is in the subhumid precipitation zone getting roughly 23 to 25 inches of annual precipitation.

Bluestem Prairie Preserve, Clay County, Minnesota. Mid-June; late vernal aspect. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 710 (Bluestem Prairie); SRM 802 (Missouri Prairie.. Plains Grassland 142.1, Bluestem "Tall-Grass" Series, 142.11 of Brown et al. (1998, p. 40). Lake Agassiz Plain 40, Glacial Lake Agassiz Basin Ecoregion 48a (White, 2020).

Four Horsemen up north- The famed "Four Horsemen of the Prairie" is an ecological appropiation of Revelation's Four Horsemen of the Aocalypse for the four dominant species of the tallgrass prairie: big bluestem, little bluestem, Indiangrass, and switchgrass. Overall these were the dominants of tallgrass grassland including that of the Agassiz Basin of the Red River Valley Region. Which plant species are dominants, co-dominants, associates etc. vary from range site to range site and, at smaller spatial scale, from local microsite to local microsite even at the climax state. Add in the seral stages of vegetation development at the same location (even at a given pasture-size scale) and from this, the derivation of range condition class and, with two determinatins of condition class, range trend over spans of time and there are a lot of changes even beyond seasonal changes like seasonal societies. Then factor in changes in plant vigor (perhaps due to weather or grazing management), in growth and maturity (phenological) stage, in soil conditions (water content, fertility, etc.), and so and things get complicated in a hurry.

Range management is an art and a craft as well as a science.

The large clumps of last year's shoots in these slides was mostly of tallgrass species including those in addition to the "big four" plus midgrasses like sideoats grama (Bouteloua curtipendula). Last season's shoots of Indiangrass and switchgrass were conspicuous in the first slide. Although much of the green color was foliage of warm-season grasses "just getting started" this far north, a lot of it was also of cool-season species especially of porcupinegrass, a spring society dominant. There were plants of the two naturalized Eurasian perennial forage grassses, Kentucky bluegrass (Poa pratensis) and smooth brome (Bromus inermis).There were also plants of the naturalized annual Japanese brome or Japanses chess (Bromus japonicus).

Forbs included purple meadow-rue, smooth scouring rush, northern bedstraw, Canada windflower, and Indian hemp. The small shrub, prairie or Arkansas rose was also "among the numbered".

Bluestem Prairie Preserve, Clay County, Minnesota.Mid-June; late vernal aspect. FRES No. 39 (Prairie Grassla.and Ecosystem). K-66 (Bluestem Prairie). SRM 710 (Bluestem Prairie); SRM 802 (Missouri Prairie.. Plains Grassland 142.1, Bluestem "Tall-Grass" Series, 142.11 of Brown et al. (1998, p. 40). Lake Agassiz Plain 40, Glacial Lake Agassiz Basin Ecoregion 48a (White, 2020).

Four Horsemen and side-kicks- Two horizon-wide views of warm-season dominated tallgrass prairie in the Red River Prairie subsection of the Red River Valley section of the Prairie Parkland province in the Minnesota Ecological Land Classification Hierachy (Minnesota Department of Natural Resources, Overall dominants were big bluestem, switchgrass, and Indiangrass. Little bluestem was one of the most widely distributed (perhaps the most widely distributed) of the dominants), but it often took a back to the other three of "the big four". In the first slide a large clump of tall dropseed (Sporobolus asper), which was conspicuous with its partly opened contracted panicle, was in left center foreground, as was one sexual shoot of tall dropseed in right immediate foreground. Sideoats grama was also present, but identifiable only by its leaves as no sexual stalks were found.

In the vernal society shown here porcupinegrass was a community dominant. Porcupiengrass appeared to be the cool-season dominant. Junegrass (Koleria cristata) was also present, but in now way was it a dominant species (except in pinpoint spots where it grew).Canada wildrye was likely present, but this rangeman could not find identifiable plants of it. There were plants of naturalized Eurasian grasses including the annual Japanese brome and the agronomic perennnials Kentucky bluegrass and smooth brome. Growth of the warm-season grass species at this point of te growing was only six to eight inches inheight.

Forb species included purple meadow-rue. northern bedstraw, Canada windflower, smooth scouring rush, golden Alexander, and Indian hemp.

A very conspicuous partial crown of false indigo bush, a woody papilionaceous legume, was in the left near foreground.

Bluestem Prairie Preserve, Clay County, Minnesota.Mid-June; late vernal aspect. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 710 (Bluestem Prairie); SRM 802 (Missouri Prairie.. Plains Grassland 142.1, Bluestem "Tall-Grass" Series, 142.11 of Brown et al. (1998, p. 40). Lake Agassiz Plain 40, Glacial Lake Agassiz Basin Ecoregion 48a (White, 2020).

Northern prairie denizens- Three "photoquadrantss" of the range plant community, the sward of the grassland, of a tallgrass prairie in the greater Red River Valley in a subhumid precipitation zone (23-25 inches annual precipitation). These views were of the vernal society (ie. spring-season, cool-season growth). Given that the Four Horsemen of the Prairies panicoid grasses that dominated this tallgrass prairie were "only just started" or barely passed spring green-up, this spring plant community differed greatly from that of the summer-autumn vegetation at peak standing crop (and grass shoot heights of four to eight feet).

Major plants in these three "photoquadrants" included porcupinegrass, the native cool-season dominant, and Junegrass, a climax indicator species. There was an unidentifiable (prebloom) Carex species.Also present in one or more "photoplots" was purple meadow-rue, goldern Alexander, northern bedstraw and a goldenrod species. Much of the plant cover and material was thatch or litter from last year's biomass most of which was that of the dominant panicoid grasses (big bluestem, Indiangrass, and switchgrass) plus tall dropseed, sideoats grama as well as cool-season porcupinegrass along with the exotic invaders Japanese brome, Kentucky bluegrass,,and smooth bromegrass.

Bluestem Prairie Preserve, Clay County, Minnesota.Mid-June; late vernal aspect. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 710 (Bluestem Prairie); SRM 802 (Missouri Prairie.. Plains Grassland 142.1, Bluestem "Tall-Grass" Series, 142.11 of Brown et al. (1998, p. 40). Lake Agassiz Plain 40, Glacial Lake Agassiz Basin Ecoregion 48a (White, 2020).

Range plants of Great Lakes (Red River Valley) tallgrass prairie

Smooth (but not like satin)- Smooth scouring-rush (Equisetum laevigatum) growing on a mesic (moderate water-holding capacity) microsite in northern tallgrass prairie. The five sexual shoots seen here were on the same plant, but at various stages of development or maturity as readily seen from the spore-bearing strobiolii.

Clay County, Minnesota. Mid-June (and still early spring).

An important--especially diagnostically--one- Porcupinegrass (Stipa spartea) growing in the virgin sod of a northern tallgrss prairie in the Glacial Lake Agasssiz Basin of the general Lake Agassiz Plain ecoregion (48A). This is part of the Red River Valley or the Western Young Drift section (Fenneman, 1938, ps. 558-588 passim) of the Central Lowlands physiographic province. Weaver and Clements (1938, p.) regarded porcupinegrass as co-dominant (with Sporobolus heterolepis) of the true prairie. Thus, preence of porcupinegrass on this bluestem-dominated prairie showed the affinity of these two major grassland communities.

Bluestem Prairie Preserve, Clay County, Minnesota. Late June, peak standing crop.

At the base- Numerous tillers of porcupinegrass arising from the rootcrown of this cool-season climax dominant prairie grass that was very much at home on a a relict (virgin sod) northern tallgrass prairie. All the Stipa species (your author followed the classic treatmemt of this genus given in Hitchcock and Chase [1951]) are cespitose (bunchgrsses) producing only tillers and not stolons or rhizomes. The specimen presented here was over two feet across at its base.

Bluestem Prairie Preserve, Clay County, Minnesota. Late June, peak standing crop.

Upper end- Sexual shoot (first slide) and panicle (second slide) of porcupinegrass at early or initiation of flowering stage. The panicle was just emerging from the boot. In fact, the photographer sort of "aassisted in the birth" by peeling back part of the boot. The boot is the general term for the topmost part of the shoot (culm and flag or uppermost leaf) that encases the inflorescence. This cool-season climax grass was completing its annual growth cycle ahead of the dominant warm-season grasses like big bluestem.

Bluestem Prairie Preserve, Clay County, Minnesota. Late June, emergence from boot-early graain phenological stage.

Out of the boot (barely)- Young panicles of porcupinegrass just emerging from the boot (as described and explained in the immediately preceding caption). The caryopses (grains) of porcupinegrass have a sharp callus that can be mechanically injurious if it works its way into thin, delicate skin such as that of sheep or in the eyes of any animal. Porcupinegrass is palatable and produces goo quality forage prior to emergence of the spearlike grain. Reference to quills of porcupines is definitely descriptve of this decreaser species..

Bluestem Prairie Preserve, Clay County, Minnesota. Late June, emergence from boot-early graain phenological stage.

One of the pretty ones- Local stand of Canada milkvetch (Astragalus canadensis) gowing in a wet depression on tallgrass prairie habitat. This was an especially nice colony or stand of robust plants. The rhizomatous feature of this showy perennial was obvious from this large, dense colony.

Cass County, North Dakota. Late June, full-bloom stage.

More of a pretty one- Closer views of Canada milkvetch growing in a wet depression on a tallgras prairie environment. A rhizomatous perennial that can bloom proficiently as seen here.

Cass County, North Dakota. Late June, full-bloom stage.

Note: "In theory" all Astragalus species have the potential to be toxic, including nitrate accumulation as well having the usual toxins of locoweedal, but Canada milkvetch is generally non-toxic. It is a fine forage species as well as fixing atmospheric nitrogen. Great addition to a wildflower garden or prairie scape. Attractive to pollinating insects. Definitely a decreaser on tallgrass prairie range.

Time for the butttercup family on the northern tallgrass prairie. Presented for viewing pleasure were two forb species of the subfamily Ranunculoideae of the buttercup family, Ranunculaceae.

In the spring wind- Canada windflower (Anemone canadensis) formed this nice litle colony on a glacial moraine tallgrass prairie in the Red River Valley or the Western Young Drift section (Fenneman, 1938, ps. 558-588 passim) of the Central Lowlands physiographic province.

Bluestem Prairie Preserve, Clay County, Minnesota. Late June, peak-bloom phenological stage.

More views of windowed blooms- Upper shoots and bloos of Canada windflower on a glacial moraine tallgrass prairie in the Red River Valley. This is a conspicuous prairie forb with eye-catching leaves and inflorescences, but it is more of an incidental plant than a dominant or asssociate species. Canada windloer is an indicator of high-quality prairie.

Bluestem Prairie Preserve, Clay County, Minnesota. Late June, peak bloom stage of maturity..

Rued purple of a northern prairie- Purple meadow-rue (Thalictrum dasycarpum) on a tallgrass prairie in the Red River Valley, Western Young Drift section (Fenneman, 1938, ps. 558-588 passim) of the Central Lowlands physiographic province. Purple meadow-rue is one of the largest Thalictrum speecies, but its purple-pigmented shoot is what is most distinctive about it. Purple meadow-rue grows across Canada andsouth to Arizona and eastward to Florida. It is one of the more widely distributed meadow-rue apecies..

Bluestem Prairie Preserve, Clay County, Minnesota. Late June, peak bloom stage of maturity..

Purple and pale cream- Upper shoot and inflorescence of purple meadow-rue that was growng on a glacial moraine tallgrass prairie in the Red River Valley of western Mainnesota.

Bluestem Prairie Preserve, Clay County, Minnesota. Late June, peak bloom stage of maturity.

Straw bedded on spring tallgrass- Northern bedstraw (Galium boreale) growing in the spring society of a northerrn tallgrass prairie in the Glacial Lake Agasssiz Basin of the general Lake Agassiz Plain in the Red River Valley. This is more or less an incidental prairie forb in the madder or coffee family (Ribiaceae)

Bluestem Prairie Preserve, Clay County, Minnesota. Late June, peak bloom stage of maturity.

Bedded blooms- Inflorescence of northern bedstraw in the spring flora of a northern tallgrass prairie in the eastern Red River Valley (Glacial Lake Agassiz Basin), Centaral Lowlands physicographic provinece. The name "bedstraw" apparently comes from the fact that the sweet-smelling shoots of this anual member of the coffee family were sometimes placed in a straw bed to give off the pleasing aroma. Yes, the seeds of Galium species were used in frontier days as a somtimes substitute of the real thing. Don't scoff: these were hard-scramble times and frontiersmen made do with what they had (Maybe we should try it just to say we did, and make us thankful for the better things we have.)

Bluestem Prairie Preserve, Clay County, Minnesota. Late June, peak bloom stage of maturity.

Not particular- Golden Alexander (Zizia aurea) growing at the edge of a second-growth red pine (Pinus resinosa) forest. While such a location suggest that this is a forest forb in actuality it is not choosy about the general habitat it chooses to call home. Golden Alexander seems to was golden on the spring prairies and all types of forest in the north woods and also in the forests and prairies of the Ozark Highlands far to the south of the northern Cental Lowlands and Lauretian Shield.

The first of these two slides presented the shoot portion of a plant while the second slide featured the basal leaves.

Lake County, Minnesota. Late June; ppeak flowering stage.

Prairie or forest- I could care less- The umbel inflorescence of golden Alexander produced in the still-spring flora of a tallgrass prairie (first and third slides) and at edge of a second-growth, red pine Great Lakes forest (second slide). This forb is not picky about its range plant community.The third slide is of a unit (one umbel) of the entire umbel inflorescence seen in the first slide.

First and third slides: Bluestem Prairie Preserve, Clay County, Minnesota. Miid-June. Second slide: Lake County, Minnesota. Obviosusly al examples at peak bloom stage of maturity.

Shrubs of Red River Valley Prairie:

Rose on the northern prairie- Prairie rose or, often, dwarf prairie rose (Rosa arkansana) Red River Valley, Western Young Drift section (Fenneman, 1938, ps. 558-588 passim) of western Minnesota.

Bluestem Prairie Preserve, Clay County, Minnesota. Late June, mid-bloom stage of maturity.

. Little rose up north- Dwarf prairie rose on a tallgrass prairiie Red River Valley, Western Young Drift section (Fenneman, 1938, ps. 558-588 passim) of western Mainnesota. This little shrub was keeping company with the cool-season grass, porcupine grass, and early growth warm-season grasses like big bluestem. R. arkansana is a widely distributed species with a species range from Maine across the eastern Prairie Provinces down in the Great Plains and Central Lowlands to south Texas.

Color of petals of prairie rose varies from a rich red to white or white "with a touch of pink" like the ones presented here.

Bluestem Prairie Preserve, Clay County, Minnesota. Late June, mid-bloom stage of maturity.

Little lady prairie shrub- A female specimen of prairie or sandbar willow (Salix interior= S. exigua subsp. interior) growing by a pothole in tallgrass prairie in the Red River Valley (Glacial Lake Agassiz Basin), Centaral Lowlands physicographic. Presence of this little tree (more like shrub) indicates a shallow water table. Members of the Salicaceae (willow and cottonwood family) are dioecious.

Bluestem Prairie Preserve, Clay County, Minnesota. Late June, peak bloom pnehological stage.

Little crown- Views of the crown of a female sandbar willow growing by a depression on a tallgrass prairie at edge of the Prairie Pothole Region. This was in the Red River Valley (Glacial Lake Agassiz Basin) of western Minnesota Taxonomy (actually, nomenclature) of sandbar willow is a garbled affair, but whatever taxon this plant is widely distributed in interior North Amereica being absent from the much of the Western Range Region and southeastern North America. Sandbar or pra;rie willow is found throughout most of the tallgrass and true prairie area of the Central Lowlands and the eastern portion of the Great Plains physiographi provinces.

Bluestem Prairie Preserve, Clay County, Minnesota. Late June, peak bloom pnehological stage.

Ecotonal Grassland- Transition of Tallgrass Prairie to True Prairie and/or Mixed Prairie

At the western edge of the Central Lowlands and the eastern margin of the Great Plains there is a physiographic section assigned to the Great Plains province and described by Fennemann (1931, ps. 25-30) as the Plains Border or, sometimes as "the break of the plains" though that has other connotations including the famed "Caprock" of Texas. Most specifically the Plains Border Physiographic section was defined and described in detail as found in Kansas by Frye and Swineford (1949).

It is in this Plains Border (the geologic/physiographic transition between Central Lowland and Great Plains physiographic provinces) which extends south into north Texas that are to be tallgrass, true, and mixed prairie grassland associations and transitions among them in a vegetational mosaic across this immense landscape.

The following section dealth with this ecotonal climax grassland by using a set of examples of these transitional range grassland communities that had developed in the Wichita Mountains section of the Central Lowlands (Fennemann, 1931, ps. 25-30) or, here again, the Plains Border section of the Great Plains physiographic province (Frye and Swineford, 1949).

Any of these examples could arbitrarily be placed in tallgrass prairie, true prairie, or mixed prairie (Clements, 1920). This professor left it up to his students to choose for themselves which, if any, of these traditional, historic, and, in this professors opinion, useful designations (Clements' associations or, in instance of tallgrass prairie, associes). Rangeland cover types reecognized by the Society fo0r Range Management (Shiflet, 1994) were given. This avoided the confusing and arbitrary designations of tallgrass, true, or mixed prairies.

128. A ~~hard~~ nearly impossible call- A consociation of little bluestem on outer margin (outskirts) of a bottomland in the Wichita Mountains of southwestern Oklahoma. Although this was a nearly single-species stand of little bluestem there were some plants of sand dropseed (Sporobolus cryptandrus) and fewer yet of sideoats grama. The associate species in this grassland vegetation was the perennial forb, Carolina nightshade (Solanum carolinense).

Question for the vegetation scientist: "Is this climax grassland tallgrass prairie or true prairie?" Yes, little bluestem is certainly a tallgrass species. It is the most xeric (least mesic) of the Four Horsemen of the Prairies; better adapted to drier soil conditions than switchgrass, big bluestem or Indiangrass (in that general order of adaptation to less moisture-favorable edaphic haibtats). So does this dense population of large plants of little bluestem (with a few midgrasses and perennial forb) delimit a tallgrass prairie? Or is this true pririe? Clements (1920, p. 122) listed little bluestem as forming one of five consociations in the true prairie association. Dodd in (Gould and Shaw, 1983, p. 346-347) regarded little bluestem as one (probably the most defining single one) species of the true prairie association. Yet, Clements (1920, p. 132) also listed little bluestem as one of eight consociations of, what in his monoclimax theory, Clements regarded a s the subclimax prairie (Andropogon associes) and what, ever since, rangemen and grassland ecologists have described as climax tallgrass prairie.

Actually the range vegetation seen here was part of the same buffalo (Bison bison) range that had the slightly different climax grassland community shown in the next (following) two slide-caption set. In fact this local vegetation (and some other local examples of this plant community) that was (were) comprised almost exclusively of little bluestem developed within an overall range plant commujnity dominated by little bluestem yet with sideoats grama as the associate to co-dominant species and with sand dropseed as a major up to local associate species. As such, this example was interpreted as a part of the general bluestem-grama cover type (SRM 709; Shiflet, 1994).

Technical note: even a quick glace at this slide and subsequent slides that followerd immediately below revealed that the current photograph had been overscanned or "bleached out" or "fadded out" in the scanning process. This was the inferior brand of scanner sold under the trade name of Epson Perfection 700 Scanner. One of this author's colleagues informed him tht it was his understanding that the Epson scanner was programmed with some random selection as to amount of light shone through (or however used) eacn slide. If this so--or, for that matter, whatever the reason--it illustrated the inferior nature of this apparatus. NEVER BUY EPSON SCANNERS!

On a positive note regarding technology, this photographer was able to salvage something out of the FUBARED mess Epson Perfection (which it is not) made of a very good slide by using Adobe PhotoShop to somewhat restore the image to the original photograph on Provia 100F film. Hurray for Adobe Photo-Shop; Hell with Perfection 700 Scanner.

Anyway, this ws one of the "puriest" (most nearly exclusive) natural stands (not a reseeded stand) of little bluestem this rangeman ever saw. It was virgin sod and the climax vegetation.

The line of trees in the background were along a small stream. Major tree species in the local forest were post oak, blackjack oak, pecan (Carya illionensis), black walnut (Juglans nigra), Arizona walnut (J. major), sugarberry (Celtis laevigata), and eastern cottonwood (Populus deltoides). Redbud (Cercis canadensis) was the main understorey shrub.

Wichita Mountains National Wildlife Refuge, Comanche County, Oklahoma. Early August; estival aspect, peak standing crop. FRES 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 709 (Bluestem-Grama). Cold Temperate Grassland 142, Plains Grassland 142.1 , Bluestem "Tall-Grass" Series 142.11 (Brown et al., 1998, p. 40). Range site: Loamy Prairie (Soil Consrvation Service, 1967). Central Great Plains Wichita Mountains 27k ecoregion (Woods et al., 2005).

129. Same range; slightly different, local range plant community- A bottomland grassland in the Wichita Mountains of southwest Oklahoma with local assemblages of grass and forb species varying from nearly single species-stands or local populations of little bluestem to dominance by little bluestem and with sideoats grama as associate species, and to co-dominance by little bluestem and sideoats grama. In assemblages of prairie plants co-dominated by little bluestem and sideoats grama, sand dropseed was the associate grass spefies. Important forbs were mostly Carolina horsenettle, wild alfalfa or slimflower scurfpea (Psoralea tenuifolia), and wooly croton (Croton capitatus). There were no woody plants in this grassland range.

An example of the "pure" stand, a consociation, of little bluestem was presented and discussed in the immediately preeding slide-caption set. At that point, the question with regard to vegetation classification centered on whether the little bluestem consociation was tallgrass or true prairie. In the grassland vegetation viewed in this set sideoats grama, a mid-grass species joined little bluestem as eith co-dominant or associate species so that this climax range plant community was clearly a mixed prairie, or (possibly, arguably) true prairie given that a little bluestem is one of five consociations of the true prairie Clements (1920, p. 122) and that little bluestem is likely the single most important--at least the most defining--plant species of the true prairie (Dodd in [Gould and Shaw, 1983, p. 346-347]).

Technical note: The Epson Perfection (It Is Not) 700 scanner used to scan these slides managed to botch up one out of five or six slides. The immediately preceding slide was an example of this imporper scanning. Fortunately, the next four slides (including the two shown here) were scanned properly (correctly) by the Epson Perfection 700 scanner.

Wichita Mountains National Wildlife Refuge, Comanche County, Oklahoma. Early August; estival aspect, peak standing crop. FRES 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 709 (Bluestem-Grama). Cold Temperate Grassland 142, Plains Grassland 142.1 , Bluestem "Tall-Grass" Series 142.11 (Brown et al., 1998, p. 40). Range site: Loamy Prairie (Soil Conservation Service, 1967). Central Great Plains Wichita Mountains 27k ecoregion (Woods et al., 2005).

130. Transition grassland- A patchwork of local grassland communities on virgin (and climax) prairie in the Wichita Mountains of southwestern Oklahoma. This example of the potential natural vegetation was in a transition zone between the Great Plains and the Central Lowlands physiographic provinces (eastern and western margins, respectively) traditionally known as the Plains Border section of the Great Plains physiographic province (Frye and Swineford, 1949) or, somewhat more specifically (or amounts to about the same thing), the Wichita Mountains section of the Central Lowlands (Fennemann, 1931, ps. 25-30). Hand in glove with a transitional physiography was the transitional (ecotonal might be a preferred ecological term) physiography, structure, and composition of grasslands within the physiographic unit.

Within the expanse of this one buffalo pasture there were various local grassland communities comprised of varying cover and density of the following grass species: little bluestem (generally the dominant species), silver bluestem (probably the overall associate species), sideoats grama, Canada wildrye, hairy grama, and prairie cordgrass (Spartina pectinata). The latter grass is a strongly rhizomatous species that often forms extensive colonies on bottomland habitats. Such was the case in local areas and, especially in an adjoining range site, in this same bison range. These local exclusive populations of prairie cordgrass constituted a consociation of the Andropogon associes (subclimax prairie association) of Clements (1920, p. 132). This Clementsian association has long been known as the tallgrass prairie (in contrast to the Clementsian true prairie). The subclimax (=tallgrass) prairie of Clenents also included consociations of the following major (local dominant or associate) species: little bluestem, silver bluestem, sideoats grama, and Canada or nodding wildrye.

From this perspective the total grassland vegetation of this one buffalo pasture could be interpreted as tallgrass prairie or, alternatively, given overall dominance of little bluestem with silver bluestem and sideoats grama as its associates, this range could be regarded as mixed prairie of the bluestem-grama rangeland cover type (SRM 709, Shiflet, 1994). Also given, however, the extensive area in a consociation of prairie cordgrass, this range could be doubly justified as tallgrass prairie. In other words, transition grassland. This explained inclusion of this climax range vegetation in the Grassland chapter entitled Tallgrass Pairie- Interior.

There was some cover and small local populations of Japanese chess or Japanese brome (Bromus japonicus), a naturalized Eurasian annual grass that is typically an invader fue to such disturbances as overgrazing, road construction, and tillage.

Important forbs in this more-or-less pristine prairie (obviously Excellent range condition class) included wholly croton, western ragweed, wild alfalfa or slimflower scurfpea and Illinois bundleflower (Desmanthus illinoensis), Baldwin ironweed (Vernonia baldwinii), and curlycup gumweed. No woody plants plagued the virgin sod of this relict grassland.

Wichita Mountains National Wildlife Refuge, Comanche County, Oklahoma. Early August; estival aspect, peak standing crop. FRES 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 709 (Bluestem-Grama). Cold Temperate Grassland 142, Plains Grassland 142.1 , Bluestem "Tall-Grass" Series 142.11 (Brown et al., 1998, p. 40). Range site: Loamy Prairie (Soil Conservation Service, 1967). Central Great Plains Wichita Mountains 27k ecoregion (Woods et al., 2005).

131. Now here's a mixture (and an vegetation classifiction "nightmare")-

Technical note: fortunately, an Epson Perfection 700 scanner did right by these two slides and the two immediately preceding sets of 35mm slides.

Wichita Mountains National Wildlife Refuge, Comanche County, Oklahoma. Early August; estival aspect, peak standing crop. FRES 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 709 (Bluestem-Grama). Cold Temperate Grassland 142, Plains Grassland 142.1 , Bluestem "Tall-Grass" Series 142.11 (Brown et al., 1998, p. 40). Range site: Loamy Prairie (Soil Conservation Service, 1967). Central Great Plains Wichita Mountains 27k ecoregion (Woods et al., 2005).

Further confusion: a seasonal or semi-wet meadow of wet prairie (not wet long enough or with soil surface inudated with water enough to be a marsh) in the same bison range that included the transition grasslands presented in the immediately preceding four slide-caption sets was a consociation of prairie cordgrass. This was obviously tallgrass prairie and not transition grassland per se as in above examples. This prairie cordgrass consociation was included at this location because it was contiguous with the climax grassland vegetation presented and discussed immediately above.

Apologies for any confusion due to this arrangement. Yes, the prairie cordgrass cover type was treated above as well as here. Again, your author let Mother Nature's arrangement on the land be the template for the organization in this chapter. Readers may take this matter up with the "laws of Nature and of Nature's God" as Thomas Jefferson so aptly put it.

132. Now this is an abrupt change and transition- In the same buffalo range having the grassland vegetation shown and described above there was this "phototransect" of a remarkable transition from tallgrass prairie to mixed prairie in the matter of a few yars. In the foreground was the perimeter of a moist to seasonally wet meadow--generally known as wet prairie--dominated by prairie cordgrass with sideoats grama as the associate species. Almost no other grass species were present, but in local habitats there were local colonies of broadleaf cattail (Typha latifolia) which, assuredly, indicated ponded water much of the year (at least growing season).

In midground to background of this image the range vegetation was mixed prairie composed of little bluestem, silver bluestem, sideoats grama, and hairy grama (in that approximate overall proportion of foliar cover). This mixed prairie had the full complement of tallgrass, midgrass, and shortgrass species as major range plants. This veteran rangeman has seldom seen such a drastic--almost "fenceline"--change in range plant communities which obviously indicated a change in range sites.

There was a narrow local ecotone or transition zone between the wet prairie of prairie cordgrass to the mixed prairie of tallgrass, midgrass, and shortgrass species.

Wichita Mountains National Wildlife Refuge, Comanche County, Oklahoma. Early August; estival aspect, peak standing crop. Two vegetational units plus a transition between them: 1) FRES 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). Prairie Cordgrass variant of SRM 710 (Bluestem Prairie). Cold Temperate Grassland 142, Plains Grassland 142.1 , Bluestem "Tall-Grass" Series 142.11 (Brown et al., 1998, p. 40). Range site: Heavy Bottomland (Soil Conservation Service, 1967). 2) FRES 38 (Plains Grassland Ecosystem). K-62 (Bluestem-Grama Prairie). SRM 709 (Bluestem-Grama). Cold Temperate Grassland 142, Plains Grassland 142.1, Mixed "Short-Grass" Series 142.13 (Brown et al., 1998, p. 40). Range Site: (Soil Conservation Service, 1967). Central Great Plains Wichita Mountains 27k ecoregion (Woods et al., 2005).

133. Wet prairie in the Wichitas- A wet prairie (at least a seasonally wet meadow) in the Wichita Mountains that was a consociation of prairie cordgrass (Spartina pectinata). The associate species--if it could with imagination be regarded as such--was sideoats grama. This was a most unusual combination of a highly mesic (almost wet or, at least, seim-hydric) tallgrass species and a mixed grass species that is well adapted to a wide range of soil moisture conditions. The principal forb was curlycup gumweed.

The wet to locally hyrdic habitats of this wet prairie was indicated by local colonies of broadleaf cattail (Typha latifolia). This wet grassland had a locally dominant grasslike plant species in broadleaf cattail.

The first of these two slides was a wide angle view of the prairie cordgrass consociation that had developed on this bottomland site. The second slide was a closer-in view of the prairie cordgrass sward. In effect, these two images constituted a "photoplot" and a "sub-photoplot" (ie. a nested arrangement of images).

This bottomland site was part of the same bison range that was featured in this section of Ecotonal Grassland in the Wichita Mountains.

134. Gently used sacrifice area- The main "water hole" for a buffalo range in the Wichita Mountains that had various grassland communities that ranged from wet prairie made up mostly of prairie cordgrass to "nearly pure" consociations of little bluestem to classic mixed prairie having tallgrass, midgrass, and shortgrass species.

The grassland vegetation presented in these two images of North American buffalo leaving a man-made lake was co-dominated by the two most mesic of the four major tallgrass species: 1) switchgrass and 2) big bluestem. Although this bottomland range site was co-dominated by these two tallgrass species there was also considerable cover of western ragweed which was a local (spot-to-spot) dominant (dominant in patches over the two tallgrass species). Other grasses included the midgrass, silver bluestem, and the shortgrass species, hairy grama. There was also considerabld cover of Canada or nodding wildrye.

Canada wildrye (a cool-season, perennial), big bluestem, and switchgrass were all decreasers on this range site while silver bluestem was an increaser and hairy grama and western ragweed were invaders. In short, there was evidence of spot overgrazing in this sacrifice area around water, but there was still greatest cover of the two warm-season decreasers which were co-dominants of this fgently used sacrifice area.

Students should use this as a textbook example of proper range management, wise and efficient use of range resources. Even the sacrifice area of this bison range was in "good shape". By the standard of range condition class even this sacrifice area was rated as Good. Superb management.

Wildlife note: the individual buffalo in this Wichita Mountains herd managed by the US Fish & Wildlife Service are "pure buffalo". They have all been blood typed to assure that all their genes (Deoxyribonucleic Acid, DNA) are Bison bison. There is no contaminating DNA from domestic cattle (Bos taurus, B. indicus) in the Wichita Mountains herd. Sadly, many supposed bison in North America still possess some cattle "blood" (DNA tracable to Bos species). This genetic contamination came about by both accident or without intention as well as through long-term (and essentially futile) attempts to develop a "breed" (a hybrid of these two bovid genera) that would be better-adapted "cattle" for harsh habitats of Great Plains range. Such efforts were made by the government such as the Canadian Department of Agriculture as well as early pioneer ranchmen such as Charles Goodnight. The Wichita Mountains bison herd serves as an outstanding source of germ plasm for preservation and production of the North American buffalo or bison.

Wichita Mountains National Wildlife Refuge, Comanche County, Oklahoma. Early August; estival aspect, peak standing crop. FRES 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 709 (Bluestem-Grama). Cold Temperate Grassland 142, Plains Grassland 142.1 , Bluestem "Tall-Grass" Series 142.11 (Brown et al., 1998, p. 40). Range site: Loamy Bottomland (Soil Consrvation Service, 1967). Central Great Plains Wichita Mountains 27k ecoregion (Woods et al., 2005).

True Prairie- short section on True Prairie inserted in Tallgrass Prairie chapter

The following example of True Prairie was placed in the Tallgrass Prairie chapter to show the closeness of these two Clementsian associations (associes in case of Tallgrass Prairie) and as a way to direct readers to the much larger chapters Tallgrass Prairie- Interior IA and Tallgrass Prairie- Interior IIB.

135. True prairie or tallgrass prairie (you choose)- In the eastern edge of the Southern High Plains immediately adjacent to the Smoky Hills province the range vegetation presented here had developed as a plant community consistenting almost exclusively of tall or meadow dropseed (Sporobolus asper var. asper= S. compositus). Very few other species were present although big bluestem made a respectible showing locally (ie. a local associate species). Local areas of heavy spot grazing (localized overgrazed patches) were populated primarily by buffalograss (Buchloe dactyloides). Otherwise, this range vegetation was almost or essentially a population of tall dropseed. Aspect was a northeast-oriented slope, but so gentle as to constitute minor impact of orientation to sun.

Successional and classification status of this range vegetation was unknown. This author did not know if this was a degraded raange on which the potential natural (climax) plant community was typical tallgrass prairie that was most llikely dominated by big bluestem (given that this species was the associate species on locally rstricted small areas [perhaps microhabitats]) or, alternatively, if this was true prairie on which tall dropseed was the climax dominant, even sole dominant such that was a tall dropseed consociation (a tall dropseed climax). Personally, the author opted for the latter, interpreting this as an example of true prairie the potential natural vegetation of which was a consociation of S. asper var. asper.

This admittedly arbitrary conclusion was based on the classic literature in the early days of American plant ecological studies. Weaver and Clements (1938, p. 318) specified that "Stipa spartea, Sporobolus asper, and S. heterolepis are the three most characteristic dominants" of the true prairie because these three species do not exist as dominants in other grassland associations. These same dominants were reported in Clements and Shelford (1939, ps. 270, 272). Tall dropseed had earlier been designated as a dominant of true prairie by Clements (1936, ps. 271, 273). Certainly, tall dropseed does occur in tallgrass prairie and mixed prairie types. It is the occurrence of tall dropseed as a dominant of true prairie that is the feature that defined the grassland shown here as being true--not tallgrass--prairie.

Presence of this range vegetation in the semiarid zone and not on mesic or postclimax habitat as, in the case of tallgrass prairie in the Nebraska Sandhills, was another convinching (to this author anyway) factor in the conclusion that this was true prairie of the climax tall dropseed form. It was observed that there were a few isolated--though small or stunted--plants of smooth brome. It was not clear how to nterpret presence of this agronomic grass in this semiarid environment. A final fact that justified this as true prairie was its proximity within a few miles of little bluestem-dominated true prairie. (This latter true prairie plant community was presented in the chapter, True Prairie within the Grassland section of Range Types of North America.)

This example of what the author interpreted as climax true prairie was included in the chapter on the more mesic and easterly tallgrass prairie to show 1) the continuity and transition of these two closely related grassland range types and 2) that classification of vegetation depends on interpretation which in turn is dependent on personal observations, experience, and, finally, biases (in turn a function of academic and experiencial factors).

Phillips County, Kansas. Late June-early estival aspect. Given published classifications this range vegetation would have the following designations FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 601 (Bluestem Prairie), generally, or, probably more specifically, SRM 710 (Bluestem Prairie). Bluestem "Tall-Grass" Series, 142.11, of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Central Great Plains- Rolling Plains and Breaks Ecoregion 27b (Chapman et al., 2001).

136. Same range in another year- Physiogonomy, structure, and species composition of a true prairie (little bluestem, dominant; tall dropseed, associate) in the more easterly portion of the Great Plains physiographic province. There were some sparsely distributed plants of Indiangrass and, in overgrazed spots, buffalograss. These were minor--almost incidental--species as, even more so, were invasive grass species such as smooth brome, Kentucky bluegrass, and Japanese brome. In field of view of these two photographs slimflower scurfpea or wild alfalfa was the most abundant forb (nodulated legume). Across this range overall, however, silverleaf scurfpea merited this designation.

These two slides of this climax range vegetation were taken two years after the photographs shown in the two immediately preceding slide-caption sets. These two "photoplots" were nested, with the subject of second slide being a smaller-scale view of the sward within the scene shown in the first slide which was a larger-scale (physiogonomic) view of this true prairie grassland.

Norton County, Kansas. Mid-June (late estival aspect). FRES No. 39 (Prairie Ecosystem). Not described by Kuchler, SRM, or Brown et al, 1998 all of whom should read the above remedial lesson (or met the old professor schooled under Dr. Albertson). Loamy Uplands range site. Central Great Plains- Rollings Plains and Breaks Ecoregion, 27b (Chapman et al., 2001).

137. Perfect example of true prairie- Truly outstanding example of a relict true prairie on rolling uplands in the Southern High Plains. This simple range plant community consisting almost exclusively of little bluestem as dominant and tall dropseed as associate species. Silverleaf scurfpea was the major forb followed by slimflower scurpea or wild alfalfa, the most common forb. There were some plants of wavyleaf thistle. In spots along cow paths and by the gates there were some plants of the naturalized Kentucky bluegrass and cheatgrass, perennial and annual (respectively) invaders from Eurasia. Otherwise this range was like it was when the Arapaho had it to themselves. Details of this range vegetation were presented below.

This range was being grazed by cow-calf pairs (black baldies) with wise-use management: correct stocking rate, proper season of use, and proper animal distribution. Ideal example of this range type in high Good to Excellent range condition class.

138. An example to educate and enlighten- Physiography of rangeland and physiogonomy (along with structure and species composition) of true prairie in eastern Great Plains. Little bluestem-dominated grassland with tall dropseed as associate species. There were also some plants of Indiangrass and, in small spots of overgrazing, buffalograss. This was not, however, anything even apporaching mixed prairie. It was classic ture prairie, the rarest of all of the major grassland associations in central North America. Japanese chess or Japanese brome, cheatgrass, smooth bromegrass, and Kentucky bluegrass,four species of naturalized Eurasian grasses, were--for all practical purposes--absent from this cow-calf range. These four invader species were present only in small disturbed areas near an adjacent tame pasture dominated by Kentucky bluegrass. They were mentioned to emphasize their diagnostic absence--for all practical purposes--from this climax grassland.

Forb species included silverleaf scurfpea, slimflower scurfpea, and wavyleaf thistle.

This landscape-scale photograph was included to present an outstanding example of true prairie. This large-scale image served to distinguish true prairie from tallgrass prairie (the bluestem-Indiangrass-switchgrass association) that is the climax (or subclimax, depending on the theory of climax one accepts) grassland association to the more mesic grasslands farther east in the Central Lowlands province.

Texas Blackland Prairie

139. Tallgrass form in Blackland Prairie of Texas-Less than 1/10 of 1% of original "Waxy Land" Prairie remains. Here in Tridens Prairie (Lamar County, Texas) is a rare upland site dominated by eastern gammagrass or, to locals, corngrass (Tripsacum dactyloides) and switchgrass. Also present arebluestems, tall dropseed (Sporobolus asper), longspike tridens (Tridensstrictus), and Florida paspalum (Paspalum floridanum) as well as forbs like rattlesnake master or snake-root eryngo (Eryngium yuccifolium) and giant coneflower (Rudbeckia maxima ). Estival aspect, June. FRES No. 39 (Prairie Ecosystem). K-68 (Blackland Prairie). SRM 717 (Little Bluestem-Indiangrass-Texas Wintergrass). Texas Blackland Prairies- Northern Blackland Prairie Ecoregion, 32a (Griffith et al., 2004).

140. Texas Blackland Prairie- The blackland or waxy land prairie has all but vanished. Less than 0.1 % of the original blackland prairie remains. The rest fell before the plow and other forms of intensive agriculture and commerce. This is a rare remnant of what was once a distinct form of tallgrass prairie. The grassland community seen here is on a hog wallow (gilgai) microrelief prairie formed by a Vertisol of montmorillonite clay that has high water-holding capacity. This edaphic habitat is dominated by eastern gamagrass, switchgrass and tall dropseed (Sporobolus asper). The conspicuous forb is yuccaleaf ernygo (Eryngium yuccifolium) but there are many other species of forbs, especially composites like giant coneflower (Rudbekia maxima), ashy sunflower (Helianthus mollis), and three goldenrod (Solidago) species. Numerous genera and families of forbs are represented as for example the bright pink corolla in the left foreground which is wild petunia (Ruellia caroliniensis).

The Nature Conservancy Tridens Prairie, Lamar County, Texas. Estival aspect, July. FRES No. 39 (Prairie Ecosytem), K-68 (Blackland Prairie), variant of SRM 717 (Little Bluestem-Indiangrass-Texas Wintergrass), Blackland range site. Texas Blackland Prairies- Northern Blackland Prairie Ecoregion, 32a (Griffith et al., 2004).

141. Graylands form of Blackland Prairie- Adjacent to the hog wallow prairie dominated by mesic tallgrass species is a droughty, acidic soil site with the unique Sylvanus or silver dropseed (Sporobolus sylveanus)-dominated prairie with Mead sedge (Carex meadii) as a local co-dominant and switchgrass, Florida paspalum (Paspalum floridanum), and rosette panicgrasses (Panicum oligosanthes and/or P. scribnerianum) are associates. Several species of less mesic forbs also occur on graylands but they contribute relatively little biomass to the total range plant community. The Nature Conservancy Tridens Prairie, Lamar County, Texas. Estival aspect, July. FRES No. 39 (Prairie Ecosystem), K-68 (Blackland Prairie), variant of SRM 717 (Little Bluestem-Indiangrass-Texas Wintergrass). Texas Blackland Prairies- Northern Blackland Prairie Ecoregion, 32a (Griffith et al., 2004).

142. Texas Blackland or Waxyland Prairie- This is another Four Horsemen tallgrass prairie but with several dropseeds including prairie dropseed and tall dropseed, longspike tridens (Tridens strictus), Florida paspalum, and cool-season grasses including Canada wildrye and Virginia wildrye (Elymus virginicus). The dominant mid-grass is sideoats grama. Texas wintergrass (Stipa leucotricha) is present but only as a spring associate or local dominant and not as a dominant of the peak standing crop community. This tallgrass community is typical of Texas prairies in that the nearly universal dominants are little bluestem and Indiangrass or little bluestem as the dominant and Indiangrass as the major associate with the other grasses varying by microsite.

There are numerous microsites on this prairie as it includes the complete sequence of blackland soils with the hog wallow (= gilgai) and mima mound relief. The latter form of microtopography occurs on clay loam or “greyland” soils. In typical prairie “fashion” there are numerous species of forbs (mostly composites). While these forbs are conspicuous they comprise relatively little of the biomass or plant cover. Dominant forbs include the yellow-flowered rough-stem or rough-leaf rosin weed (Silphium radula) and compassplant (S. laciniatum) and the annual American basketflower or American knapweed (Centaurea americana).

The prairie seen here is the least mesic upland form of blackland (in contrast to the clay soil upland prairie represented by the preceding slide of Tridens Prairie). One of the soil series present in isolated patches is the classic Houston black clay. Many of the depressions, especially those of Houston black clay, host the the unusual prairie cray(w)fish (Procambarus gracilis) which sinks its shafts down to the wet soil layers, or perhaps to the surface aquifer beneath the virgin sod. This indicates that these prairie soils are more or less permanently wet. Though this is an upland prairie, it is a wet prairie with a hydric water regime (not a marsh of inundated soil with standing surface water but the next thing to it).

Trees along the edge of the photograph are part of a gallary forest along a prairie creek. Tree species at edge of prairie include cedar elm (Ulmus crassifolia), hedge apple or bois-d-arc (Maclura pomifera), and honey locust (Gleditsia triacanthos).

The Nature Conservancy Clymer Prairie, Hunt County, Texas. Estival aspect, July.FRES No. 39 (Prairie Ecosystem), K-68 (Blackland Prairie), SRM 717 (Little Bluestem-Indiangrass-Texas Wintergrass) or variant thereof (the Blackland Prairie is more an extension of the tallgrass [Andropogon-Sorgastrum] prairie than the true [Sporobolus-Stipa] prairie as was incorrectly stated under 717 but it does contain dominant species of both). Clay Loam and trace of Loamy Prairie range sites. Texas Blackland Prairies- Northern Blackland Prairies Ecoregion, 32a (Griffith et al.l, 2004).

143. Detail of the tallgrass sward of Texas Blackland Prairie- This is a little bluestem-Indiangrass dominated tallgrass stand (the tallest and grey-green grass shoots are of Indiangrass) but associated species do contribute appreciable biomass and cover. Gramineae associates include Canada or nodding wildrye, Virginia wildrye, prairie dropseed, tall or meadow dropseed, and longspike tridens. Forb associates include rough-stem rosinweed, compassplant, American basketflower, and Maxmillian sunflower (composites are far more important than all other forb families combined). It should be specified that the dominant cool-season grasses are the wildryes and not Texas wintergrass. The SRM cover type title and description of Little Bluestem-Indiangrass-Texas Wintergrass would be Little Bluestem-Indiangrass-Wildrye for this variant, and this is a tallgrass prairie not a true prairie form.

The Nature Conservancy Clymer Prairie, Hunt County, Texas. Estival aspect, July. FRES No. 39 (Prairie Ecosystem), K-68 (Blackland Prairie), SRM 717 (Little Bluestem-Indiangrass-Texas Wintergrass) or variant thereof. Clay Loam range site.Texas Blackland Prairies- Northern Blackland Prairie Ecoregion, 32a (Griffith et al., 2004).

144. American basketflower or American knapweed (Centaurea americana)- The niche of this native annual is interesting given that annuals, including this species, usually are more common on disturbed areas. Here on virgin sod of a tallgrass prairie dominated almost exclusively by perennial grasses and forbs this annual composite is obviously plentiful and reproductively successful. This illustrates the great biodiversity possible on natural grassland communities. Clymer Prairie, Hunt Texas, July.

145. Island of Blackland or Waxyland Prairie in Texas Pineywoods- An isolated remnant tract of the “greyland” form of Blackland Prairie with pronounced "hog wallow" microrelief. This type of Waxyland Prairie develops on clay loam soils of the Vertisol order. This remarkable meadow was in the area where the western perimeter of Texas Pineywoods and the easternmost part of the Post Oak Savanna come together rather than in the Blackland Prairie Vegetational Area of Texas (Gould, 1962). A long "lineage" of private landowners had obviously appreciated the value of this native grassland vegetation for a hay meadow which, based on its gilgai microtopography, had also apparently never been plowed. (Gilgai features can reform on plowed Vertisols, but the time frame for such natural restoration is decades [Diggs et al., 2006, p. 63) and it is highly unlikely that land having such remarkable "hog wallow" microtopography had ever had a bloody plow in it.)

Unfortunately the virgin soil of this east Texas meadow did not correspond to virgin vegetation. Bahiagrass (Paspalum notatum), an introduced agronomic grass (from South America), had invaded this climax tallgrass prairie so that little bluestem and Indiangrass, the dominant decreasers, had been largely displaced by the naturalized (and often weedy) bahiagrass. While there were local spots in this meadow where little bluestem, Indiangrass, tall dropseed, upland switchgrass, and the native midgrass, sideoats grama, were dominant (examples shown below) such local areas constituted a small proportion of this potential tallgrass prairie. The large composite forb, giant coneflower (Rudbeckia maxima), was also locally dominant and served as a "botanical fossil" as to what once was. One-flower flat (= umbrella) sedge (Cyperus retroflexus) was also locally abundant.

Diggs et al. (1999, ps. 37, 41-42) cited work which estimated that that somewhere between 98 to 99.9 % of the original (just before European settlement) Blackland Prairie had been destroyed by plowing and that much, if not most, of the fractional remainder had been severly disturbed by overgrazing, overmowing, invasion by noxious exotic plants, and herbicidal treatment (this latter largely eliminates forbs, many of which add nutritious dry matter to prairie hay and diets of grazing animals).

Such combination of these conditions had drastically altered the species composition and structure of the (former) tallgrass meadow presented here. This meadow was most likely subjected to some improper haying practices such as mowing too frequently, too closely (too low a stubble height), and too late in the growing season. This last form of mismanagement or abuse constituted improper season of use. Too-late haying prevents replinishment of food reserves in roots and rootcrowns for maintenance (respiration) during winter dormancy thereby resulting in winter-kill. Invasion by bahiagrass exacerbated degratation of this meadow's plant community. Such invasion was probably facilitated by improper mowing and, perhaps, periodic grazing by cattle (meadow was fenced, but the author did not observe any cattle dung).

Seedlings of common persimmon (Diospyros virginiana) were present in locally large numbers, and mechanical mowing (albeit it imporper in some aspects) had kept this native, woody species from further invading the virgin sod. Natural fire (or prescribed burning) which also would have eliminated and prevented invasion by persimmon, but in lieu of this natural process mowing had performed this beneficial service. In fact, without mowing and in absence of fire persimmon would have converted much of this meadow into a persimmon thicket. Seed source of persimmon was trees in the fencerow perimeter of this local tallgrass prairie.

Anderson County, Texas. October. FRES No. 39 (Prairie Ecosystem), K-68 (Blackland Prairie), SRM 717 (Little Bluestem-Indiangrass-Texas Wintergrass) or variant thereof. Clay Loam range site. East Central Texas Plains- Northern Post Oak Savannah Ecoregion 33a (Griffith et al.l, 2004).

146. Hog wallers and other holes- Examples of "hog wallow" depressions of the gilgai microtopography (microvalley and microridge) on an island of Texas Blackland Prairie at western perimeter of Pineywoods. The first slide showed the textbook "hog wallow" microrelief while the second slide presented the "roof ridge" feature (to the left of and making up one side of another "how wallow". Gilai on Blackland Prairie forms on Vertisols die to shrinking and swelling of these soils under varying regimes of precipitation and temperature. Brief details of gilgai and mima mound microrelief are readily available in two excellent flora: 1) Diggs et al., 1999, p.39) and 2) Diggs et al. (2006, ps. 61-65).

Little bluestem was well-represented in the range vegetation shown in both photographs, but the less conspicuous bahiagrass was better represented. Bahiagrass is a major weed is this area, even on domestic permanent pastures where it outcompetes and displaces more productive cultivars of more desirable tame grass species such as bermudagrass (Cynodon dactylodon). A few plants of Indiangrass and even fewer of upland switchgrass gave evidence as to composition of the climax plant community for this range site. Giant coneflower was abundant enough that one could not take photographs of the meadow without including some of this dominant (and about the only) forb. One-flower flat (= umbrella) sedge was also present and sometimes locally abundant. Its successional status was not known.

147. Tallgrass and big composites- Area of a Blackland Prairie within Texas Pineywoods on which little bluestem was the local dominant grass with sparse cover of Indiangrass and upland switchgrass, associate grass species, and giant coneflower (Rudbeckia maxima) varied locally from overall dominant to associate species. Tall dropseed and sideoats grama were present in small propostions, but were also grasses that likely were component species of the climax range plant community, the potential natural vegetation before being greatly altered by white man. Although this delightful little prairie had been wisely (perhaps miraculously) spared from the plow and properly saved as a hay meadow, it had been mowed improperly (probably too frequently, too closely, and with wrong timing) for so long that it had been degraded to the point that bahiagrass had "taken over" much of the meadow.

This was a relict spot within the relict prairie on which native species had made their last stand.

148. On the edges- Border of a Blackland Prairie hay meadow in an east Texas in a "border land" between the Post Oak Savanna and Pineywoods (ie. extreme eastern and far-western perimeters of these respecitve vegetational areas). The two large trees in the background perimeter of this meadow were loblolly pine (Pinus taeda), left, and southern red oak (Quercus falcata), right. These trees (and those visible around border of the meadow in photographs presented above) stood as evidence of this small parcel of natural grassland that had developed and persisted as a "floristic island" in a transition area (a fairly broad ecotone) dominated by trees. From the perspective of Landscape Ecology this meadow of tallgrass prairie was a patch within a matrix of forest and savannah.

149. Giant coneflower (Rudbeckia maxima)- The dominant forb on a hay meadow of an isolated patch of Blackland Prairie in the transition of Post Oak Savanna and Pineywoods in east Texas. Giant coneflower is one of the most conspicuous and distinctive of the prairie forbs in the prairies and savannahs of eastern and central Texas. The large pale-blue leaves and general large size of individual plants of giant coneflower make it a good species for neophyte prairie fans and other beginners to learn. The plants shown in these and preceding photographs were all regrowth following recent haying.

Anderson County, Texas. October; late-season flowering stage in plant regrowth.

150. One-flower flat (= umbrella) sedge (Cyperus retroflexus)- This was a common species on the degraded Blackland Prairie featured here. It is a widely distributed Cyperus species throughout much of central southern North America (Alabama to Arizona. One-flower flat sedge occurs from east to west Texas. Successional status and response of this species to disturbance (decreaser, increaser, or invader) was not known, but it was likely not a member of the climax plant community (at least not at any substantial cover or abundance).

Anderson County, Texas. October.

Prairie Peninsula: Prairie-Deciduous Forest Ecotone (hardwoods-tallgrass savanna)

One of the largest ecotones in North America is the vast transition between eastern deciduous forest formation and the immense humid-to-subhumid tallgrass and true prairies portion of the central grasslands. This ecotone manifests itself primarily as a savanna of various oak and hickory species with an herbaceous understorey of tallgrass and midgrass species plus attendant forb and shrub species. In one of the classic ecological monographs of North American vegetation Transeau (1935) mapped this transition of climax plant communities and labeled the huge savanna the Prairie Peninsula.

Subsequent to establishment of the Prairie Peninsula as a more northern manifestation of the tension zone between eastern deciduous forest and central prairie (Transeau, 1935) later workers interpreted similar climax vegetation to the south of the mapped Prairie Peninsula as islands or outliers the savanna, the ecotone, first described by Transeau (1935). The Cross Timbers and much of the Ozark Plateau are now regarded by range and vegetation scientists as part of the greater Prairie Peninsula. These latter vegetational-physiogrphic-geologic areas or units were treated in a separate chapter herein as Tallgrass Savanna under the Grasslands section. This savanna range vegetation was interpreted by the current author as a form of the general central grassland formation more than deciduous forest formation (a rangeman's bias perhaps) yet treated as a distinct major unit of North American vegetation.

A brief sample of the Prairie Peninsula within the original region mapped and described by Transean(1935) was included in the present Tallgrass Prairie chapter to provide continuity in treatment and connections among coverage of the various range cover types.

153. A taste of the Prairie Peninsula- Border of natural vegetation at one of infinite (at one time) "confluences", meetings, or minglings of tallgrass prairie and eastern deciduous forest communities. Important prairie species of the grassland included big bluestem, eastern gamagrass, prairie cordgrass, prairie dropseed, switchgrass, Indiangrass, and giant goldenrod . Woody vegetation was a gallary forest along Cub Creek comprised of numerous tree species including green ash, red mulberry, black walnut (Juglans nigra), silver maple (Acer saccharinum), honey locust (Gleditsia triacanthos), and western hackberry.

Homestead National Monument, Gage County, Nebraska. July, estival aspect. FRES No.39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 601 (Bluestem Prairie). Plains Grassland 142.1, Bluestem "Tall-Grass" Series, 142.11 of Brown et al. (1998, p. 40). Central Great Plains- Rainwater Basin Plains Ecoregion, 27f (Chapman et al., 2001).

154. Landscape-scale view of the western Prairie Peninsula- Near the western extremity of the Ozark Plateau and beginning of eastern Cherokee Prairie (Central Lowlands physiographic province) the famed Prairie Peninsula of tallgrass prairie and oak-hickory forest have been in slow but perpetual battle for supremacy of the land. Overall dominant plant across this indescribably beautiful "prairiescape" was big bluestem. Local dominant was upland switchgrass, accompanied by Indiangrass, prairie cordgrass, little bluestem, and several dropseed species. Dominant forb was the large composite, compassplant (Silphium lacinatum).

Oak-hickory forests typical of those in the adjoining Ozark Highlands (Springfield Plateau) had developed on some north slopes and moister valleys of this landscape.

Prairie State Park, Barton County, Missouri. Late July, peak standing crop overall with switchgrass in full-bloom, big bluestem just beginning to elongate shoots, and compassplant at peak bloom. FRES No.39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). K-73 (Mosaic of K-66 [Bluestem Prairie] and K-91 [Oak-Hickory Forest]. SRM 802 (Missouri Prairie), essentially by definition, but SRM 601 (Bluestem Prairie) is likely a better fit. Plains Grassland 142.1, Bluestem "Tall-Grass" Series, 142.11 of Brown et al. (1998, p. 40). Central Irregular Plains- Cherokee Plains Ecoregion, 40d (Chapman et al, 2002).

155. A perfect prairie in the Prairie Peninsula- Three views of a hay meadow that was a consociation--a nearly "pure" stand as if a field crop--of big bluestem. There were almost no other plant species prsent other than occasional plants of forbs such as black-eyed susan (Rudbeckia hirta) seen here in all three "photplots" and yuccaleaf eryngo (Eryngium yuccifolium) as seen in the next slide/caption set.

This meadow had been hayed for decades and was in the most nearly "mint" (pristine) condition of any meadow observed in this general area of the western Ozark Uplift (Springfield Plateau). The landscape feature of the Prairie Peninsula was presented in these views where oak-hickory forests surrounding this tallgrass prairie were visible in the distant background.

This was a priceless relict tract of the Burkhart Prairie located sporadically over an area extending five to eight miles north of Seneca, Missouri.

Newton County, Missouri. Late July; early shoot elongation in big bluestem, immediate pre-bloom stage of phenology with a few tillers having progressed to shoot elongation atate of morphology. FRES No.39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). K-73 (Mosaic of K-66 [Bluestem Prairie] and K-91 [Oak-Hickory Forest]. SRM 802 (Missouri Prairie), essentially by definition, but SRM 601 (Bluestem Prairie) is likely a better fit. Plains Grassland 142.1, Bluestem "Tall-Grass" Series, 142.11 of Brown et al. (1998, p. 40). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Chapman et al, 2002).

156. Closer-in views of a perfect prairie- Hay meadow in western Springfield Plateau that even after haying for decades was a consociation of big bluestem so "pure" (homogenous) in species composition as be the closest ideal to pristine tallgrass prairie as any similar other example found in this area by this author, who incidentally grew up in this locality.

The first slide presented a local spot in which big bluestem had recently lodged (probably temporarily) as a result of heavy rain and high winds from a thunderstorm. Lodging is the condition of herbaceous cover or foliage being flattened (and, often, entangled) by action of storm or, sometimes, just gravity on extremely tall, heavy shoots.

The second slide was a "photoplot" of this climax vegetation in which a large specimen of yuccaleaf eryngo, a forb of the carrot or parsely family (Umbelliferae), made a "rare appearance" in a homogenous stand of big bluestem.

157. Ubelliferous forb on a perfect prairie- Details of sexual (flowering) shoots of yuccaleaf eryngo (Eryngium yuccifolium) growing in a "solid" stand of big bluestem on a remnant of tallgrass prairie (one of the scattered remains of Burkehart Prairie) in the western Springfield Plateau in southwest Missouri.

This native, perennial prairie forb is a member of the Umbelliferae or carrot family.

Technical note: the over-exposed condition of the first two slides was due to improper maintenance of the Nikkor lens that was used. It seems that several years prior to these photographs, the preventative maintenance (so-called) was performed by a supposedly knowledgable and skilled technician. Unforttunately, said technicial was incompetent had used grey grease instead of proper lens lubricant. This was unknown to the photographer. Eventually the grease hardened such that the shutter had a delayed response to manual operation. Adobe PhotoShop was used to salvage something of the image.

Newton County, Missouri. Late July; peak standing crop, immediate post-flowering stage.

158. Prairie Peninsula at union of Ozark Plateau and Cherokee Prairie- Well-maintained prairie hay meadow situated within oak-hickory forest on the ancient hills of the Springfield Plateau portion of the Ozark Mountains. Big bluestem, Indiangrass, upland switchgrass, prairie dropseed, purpletop, Canada wildrye, Virginia wildrye, and common or whole-leaf rosinweed (Silphium integrifolium) were major species in roughly that order. One common grasslike plant was tall or or littlehead nut-sedge (Scleria oligantha).

The tracts of oak-hickory forest that were "inter-woven" with tallgrass prairie were dominated--alternatively-- by post oak, blackjack oak, black oak (Quercus velutina), chinquapin oak, and/or black hickory (Carya texana), in locally varying combinations and with such associates as western hackberry, green elm, black walnut, American elm, honey locust, common persimmon (Diospyros virginiana), and sassafras (Sassafras albidum). Redbud (Cercis canadensis) and flowering dogwood (Cornus florida) dominated an upper shrub layer wereas blackberry made up a lower shrub layer. Grass species of the adjoining tallgrass prairie formed an herbaceous understorey. The wildryes and purpletop were typical major grasses. In more open oak-hickory stands big bluestem was dominant as reflected by the local common name of "timbergrass".

Ottawa County, Oklahoma. July, nearing peak standing crop. FRES No.39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). K-73 (Mosaic of K-66 [Bluestem Prairie] and K-91 [Oak-Hickory Forest]. SRM 710 (Bluestem Prairie). Plains Grassland 142.1, Bluestem "Tall-Grass" Series, 142.11 of Brown et al. (1998, p. 40). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al. (2005).

159. Sward of the western Prairie Peninsula- Another (and more detailed) view of the species-rich prairie hay meadow introduced above that was at western edge of the Springfield Plateau and beginning of Cherokee Prairie. In this view ashy sunflower (Helianthus mollis) was conspicuous along with lots of prairie blazing star or prairie gayfeather (Liatris pycnostachya).The Four Horsemen of the Prairies were in firm command, but late season haying (late July-early August) had reduced cover of these climax dominants. Increasers like purpletop had replaced some of the original cover of tallgrass decreasers, especially big bluestem (the natural dominant and defining species of this tallgrass prairie vegetation).

Species of neighboring oak-hickory Ozark forests were provided in the immediately preceding caption.

160. Recovery of an Ozark Plateau prairie- This is a portion of the prairie hay meadow presented and described in the two immediately preceding slide/caption units after three years of not making hay after haying for at least three/fourths of a century (probably longer). Most of the taller, larger grass shoots in this image were upland switchgrass. Big bluestem was also but at lower foliar cover. The main reason that big bluestem was not conspicuous in this image whereas switchgrass was is that big bluestem is a short-shoot tallgrass that does not elongate its sexual (flowering) shoots until late in the warm growing season while switchgrass is a long-shoot tallgrass that elongates shoots that development into sexual shoots comparatively early in the warm growing season.

Switchgrass and big bluestem had declined almost to the point of being absent from this meadow due largely to the practice--followed for over fifty years--of harvesting for hay in late summer or early fall. Mowing in the latter stage of the growing season--presumedly to get maximum yield of dry matter (as measured by the greatest number of hay bales--is about the worst possible time to harvest tallgrass species like big bluestem and switchgrass, the potential climax dominants on this range site. Tallgrass species mowed late in the growing season but just prior to the first killing freeze will produce some regowth at tremendous expense to root reserves of total available carbohydrate. Under these conditions late-mowed plants may winter kill because they will not have adequate root/rootcrown reserves to survive winter dormancy.

Three years of non-mowing had permitted switchgrass (and big bluestem) to recover to the extent shown here. An interesting aside fact was that the owners of this ancient hay meadow no longer neeeded the hay because they had sold their cattle (due to bad fences and no inclination to rebuild them). This is just one more example of how cultural factors, including family affairs, directly impact the land and its range vegetation.

In the short term, the dominant switchgrass and big bluestem had benefitted from cessation of late-season mowing. In the longer term, however, cessation of all mowing (and it was never burnt) of this tallgrass prairie resulted in other vegetational developments. Click for the nexty slide...

Ottawa County, Oklahoma. Mid-July; early estival aspect. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). K-73 (Mosaic of K-66 [Bluestem Prairie] and K-91 [Oak-Hickory Forest]. SRM 710 (Bluestem Prairie). Plains Grassland 142.1, Bluestem "Tall-Grass" Series, 142.11 of Brown et al. (1998, p. 40). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al. (2005).

161. Ozark Prairie- Another view (in another year) of the same hay meadow described in the three preceding photographs. This photgraph attempted to portray the mima mound microtopography that is characteristic of virgin sod of tallgrass prairie in the Ozark Plateau. The taller, greener foliage is switchgrass which was growing at base of and up on (all over) mima mounds. Major forbs were giant ironweed (Vernonia crinita) such as the fine specimen in lower left corner, ashy sunflower, yuccaleaf rattlesnake master, prairie gayfeather or prairie blazingstar, whole-leaf rosinweed, whorled milkweed (Asclepias verticillata), foxglove beardtongue (Penstemon digitalis), Culver's root (Veronicastrum virginicum), and fringed poppy mallow or winecup (Callirhoe digitata var. digitata).

Ottawa County, Oklahoma. Late July, nearing peak standing crop. FRES No.39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). K-73 (Mosaic of K-66 [Bluestem Prairie] and K-91 [Oak-Hickory Forest]. SRM 710 (Bluestem Prairie). Plains Grassland 142.1, Bluestem "Tall-Grass" Series, 142.11 of Brown et al. (1998, p. 40). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al. (2005).

162. Some Ozark prairie forbs- Local assemblage of native plants on a tallgrass prairie in the western Ozark Plateau. This partial species "photolist" was taken on the same meadow used for prairie hay production that was shown in the immediately preceding photographs. Students should take a little "phot-pop quiz" and see how many species (at least genera or, maybe, families) they can identify. STOP and write down your answers. Now you may proceed. Answers (left to right): whorled milkweed (Asclepias verticillata), prairie gayfeather or blazingstar(Liatris pychostachya), and yuccaleaf rattlesnake master (Eryngium yuccifolium) in Asclepediaceae, Compositae (Asteraceae), and Umbeliferae (Apiaceae), respectively. Actually, there was a fourth forb (and fourth plant family)--though at fruit not flower stage--in center far midground: foxglove beardtongue (Penstemon digitalis) of Scrophulariaceae.

Whorled milkweed was sheading fruit (timing can be almost everything in plant photography).

Other forbs on this hay meadow were listed in the preceding captions.

Lesson: range plant communities labeled as grassland (of the grassland biome) were so designated based on dominance by grass species of the potential natural (climax) vegetation. Such designations were (are) determined based on plant family, category (group), or life form of the dominant species where dominance is defined as to relative proportions (species composition) of the plant community based mostly on foliar or basal cover (hence dominance types= cover types). Other, and usually secondary, population parameters (biostatistics) in determination of dominance sometimes include density, frequency, general abundance, etc. This is the case for biomes and lower hierarchial units therein (associations, faciations, ... all the way down to range site). Hence: shrubland (includiing deserts and chaparral), forests and woodlands, even forbland. There are exceptions to this code or format such as alpine or tundra.

Dominance does not refer to number of species in this or that plant family, not which family or group of range/forest plants (grasses, grasslike plants, forbs, shrubs, and/or trees) has the most (greatest number) of species in the community. Rather, domiance refers to which species, family, or plant group occupies the most "volume" (= has the most cover, foliar or basal) such that they "control the high ground" of the canopy layer. On grasslands there are almost always far more species of forbs than of grasses, but grasses dominate the canopy layer (have the greatest cover) and produce the vast bulk of biomass, at least by end of growing season or cumulatively throughout the growing season.

On the virgin sod of the prairie hay meadow featured there were about as many (maybe even more) species of composites than of grasses. Throw in legumes, plus members of the snapdragon, carrot or parsley, milkweed, mallow, and other forb families and grasses were outnumbered many times over--as to numbers of species. Again, however, it is cover, biomass, and related measures of dominance not numbers of species--not biological diversity--that determines the cover or dominance types of native vegetation. One species (big bluestem in the climax vegetation described here) will commonly have more cover or greater standing crop than all other species combined. Such single-species dominance (cover) types are consociations (of the Clementsian association). More commonly there will be two or three co-dominant species. The common or scientific names of the dominants provide the title of the dominance (cover) type as, in this instance, the rangeland cover type entitled and described (Shiflet, 1994) as Bluestem Prairie (SRM 601 and/or 710).

Incidentially, the main nonforb herb (the local dominant) on many microsites of this meadow was the grasslike plant species, tall or littlehead nut-sedge (Scleria oligantha) of Cyperaceae. Sometimes species of grasslike plants dominate grassland although this is more commonly the case for marshes, mountain meadows, and related types like alpine vegetation. In this example, local dominance by nut-sedge was due to range retrogression. Littlehead nut-sedge was an ecological invader.

163. Natural grass-legume pasture mix- Big bluestem and catclaw sensitivebriar growing side-by-side on an "island" of tallgrass prairie situated within a oak-hickory savanna-forest dominated by black oak (Quercus velutina). Big bluestem was the dominant and Indiangrass was the associate on the tallgrass prairie. From the perspective of Landscape Ecology the smaller natural parcels of prairie can be viewed as patches within the matrix of oak-hickory forest and savanna. Both savanna and forest forms of the oak-hickory cover type had two or more herbaceous layers the taller of which was dominated by big bluestem and various native legumes.

Ottawa County, Oklahoma. Late May (late vernal society). FRES No.39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). K-73 (Mosaic of K-66 [Bluestem Prairie] and K-91 [Oak-Hickory Forest]. SRM 710 (Bluestem Prairie). Plains Grassland 142.1, Bluestem "Tall-Grass" Series, 142.11 of Brown et al. (1998, p. 40). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al. (2005).

164. Species diversity on an Ozark prairie- Sward of tallgrass prairie that had been degraded by decades of mowing for prairie hay in late summer to early autumn. Big bluestem had been largely replaced by tall or or littlehead nut-sedge (Scleria oligantha). In certain microsites forbs were a major component of this abused grassland vegetation. This was especially the case for those forb species that completed their annual growth cycles before destructive late-season hay harvest (ie. such species were largely unaffected by this late-date defoliation). Most of these spring-flowering forbs were climax or at, least, advanced seral species.A species-rich sample of these early growing season forbs was presented in these two "photoquadrants".

Forbs included catclaw sensitivebriar, ashy sunflower (Helianthus mollis), ground plum (Astragalus crassicarpus var. trichocalyx= A. mexicanus var. trichocalyx), five-finger cinquefoil (Potentilla canadensis var. villosissima), Sampson's snakeroot (Psoralea psoralioides var. eglandulosa) and, the centerpiece of God's own bounteous bouquet,.ragged orchid (Habenaria lacera var. lacera). The main graminoid was littlehead or tall nut-sedge. The most common grass was winter bentgrass (Agrostis hyemalis). All of these species were visible in the first slide. Species details were more prominent in the second slide which was a closer-in view yet with fewer plants of the different species.

165. Another sample of species richness in the Ozarks- A third "photoplot" of the sward of a tallgrass prairie degraded by late summer to early autumn haying (the same meadow as shown in the preceding two-slide set). Range plant species in this plot included Junegrass (Koleria cristata), Sampson's snakeroot, littlehead or tall nut-sedge, ground plum, and catclaw sensitivebriar. These were all species that flowered and set fruit in late-spring to early summer.

Range plant species--including forbs, grasses, and grasslike plants--that flower in spring to early summer (ie. those that completed their annual cycle far in advance of destructive late-season mowing) were favored over (at a competitive advantage relative to) late summer- or autumn-flowering species. These latter tend to be the larger, dominant, climax species (decreasers such big bluestem, switchgrass, prairie dropseed) under proper management. Cool-season grasses, including natives like Junegrass and winter bentgrass were largely unaffected by late summer or early autumn hay harvest because by this point in time cool-season species had long been dormant. By contrast, the late-date haying coincided with defoliation at advanced phenological stages and when little of the warm-growing season remained for native warm-season climax (decreaser) grasses. It was far too late in the annual growing cycle for these dominant and associate species of tallgrass prairie to replinish storage reserves in rootcrowns and roots which is essential for survival during winter dormancy. This is a textbook example for one of the Cardinal Principles of Range Managment: Proper Season of Use.

Prairie hay should be in the bale by no later than mid-July in northern Oklahoma and southern Kansas. Late haying was improper management, and both stupid and ignorant as any observant hay-maker should--with years of experience--be astute enough to figure this out. Such improper management did favor some interesting species even if not the potential natural dominants and the most productive and palatable forage species.Such a pattern of defoliation might well have favored Junegrass and permitted greter density, cover, etc. of this midgrass species than if it had to compete with tallgrass species (even in early parts of the warm-growing season).

166. Most species diversity yet- Rainbow of species on a prairie hay meadow in the Springfield Plateau. The first slide included an array of range plants that ranged through grasses, forbs, and even a shrub. Forbs included three legumes all of which were decreaser species: catclaw sensitivebriar, leadplant (Amorpha canescens), and wild or blue false indigo (Baptisia australis var. minor). Grass species ran the gamet from decreasers (big bluestem and switchgrass) to an invader (broomsedge bluestem). The shrub, New Jersey tea (Ceanothus americanus), was a decreaser. Quite a coalition, and one of mostly climax species.

These "photoplots" were from a more protected area of a hay meadow that, like the meadow from which the "photosamples" shared immediately above were from, had been degraded through decades of late-summer haying. Big bluestem and switchgrass were the potential natural (climax) dominants, but decreaser species were quite limited and both tallgrass prairie meadows were in a state of deterioration (retrogression) so as to rate in only Fair range condition class. Range vegetation shown here was in a back corner of the hay meadow where local microtopography of mima mounds afforded more protection to prairie plants.

Newton County, Missouri. Late May (late vernal society). FRES No.39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). K-73 (Mosaic of K-66 [Bluestem Prairie] and K-91 [Oak-Hickory Forest]. SRM 710 (Bluestem Prairie). Plains Grassland 142.1, Bluestem "Tall-Grass" Series, 142.11 of Brown et al. (1998, p. 40). Ozark Highlands- Springfield Plateau Ecoregion, 39a (Woods et al. (2005).

Wet Prairies and Marshes of Tallgrass Prairie

Within the extensive tallgrass prairie region there are range types and range sites of wet prairie and marsh. Marsh is commonly regarded that kind or form (type) of wetland dominated by herbaceous (vs. woody) vegetation. Marsh was defined by Mitsch and Gosselink (2007, p. 32) as "a frequently or continually inundated wetland characterized by emergent herbaceous vegetation, adapted to saturated soil conditions". More specifically and in common usage among rangemen marshes are interpreted as wetlands dominated by grasslike plants, especially those of Cyperaceae, Juncaceae, and Typhaceae, rather than grasses (Gramineae) although grasses are frequently major marsh species. Thus, by converntion marshes are distinct from grasslands, including wet grasslands. The two most common types (kinds or forms) of wet grasslands are:1) wet prairies, especially, and 2) wet meadows, secondarily (when dominant plants are grass species).

Mitsch and Gosselink (2007, p. 33) defined wet meadow as "grassland with waterlogged soil near the surface but without standing water for most of the year" whereas wet prairie was described as "similar to a marsh, but with water levels usually intermediate between a marsh and a wet meadow". In other words, these authors defined units of natural wetland vegetation based on soil water parameters (conditions). In this chapter on tallgrass prairie and throughout Range Types the current author definded and distinguished among such herbaceous wetlands as marshes, wet prairies, and wet meadows primarily as to vegetation (range plant communities) and secondly based on edaphic water criteria.

A complication and source of confusion in description of range vegetation naturally arises when marshes such as those dominated by rushes, sedges, bulrushes, cattails, etc. occur as units of natural range plant communities within the general, surrounding tallgrass prairie region, especially when there are contiguous range types that form continua of native vegetation ranging from dry upland to mesic tallgrass prairie, wet tallgrass prairie, and marshes that are dominated (if not comprised completely) of grasslike plant species.

Therefore, for consistently in treatment of range vegetation, marshes and meadows were not regarded herein as grasslands or grassland types. However, for some degree of continuity and to facilitate locating of range types herein as well as understanding relations among range types some examples of marshes--along with wet prairies--were included immediately below to avoid confusion and represent the continuum of mesic prairie, wet prairie, and marsh.

For these same reasons, marshes, lake vegetation, and related wetlands in the Nebraska Sandhills were also included with the postclimax tallgrass prairies of semiarid areas that was included later in this chapter. Otherwise, marshes were treated separately in Range Types of North America under the chapter, Meadows and Related Marshes.

Continuum of Wet Prairie to Marsh

A mosaic of wetlands ranging from wet prairie to river floodplain to marsh exist in the immediate vicinity of the Salt Fork of the Arkansas River in northern Oklahoma and southern Kansas. This natural "patchwork" of various grasslands and marshes provided a good example herbaceous wetlands within the tallgrass prairie region. Marshes are not grasslands strictly speaking or in precise usage, but in the example provided here (and it was a typical situation over formerly large areas in the once-vast grassland domain of central North America) marshes and wet prairie formed an intricate, inter-related herbaceous vegetation at both landscape and ecosystem levels. These two biomes were interconnected by processes at landscape-scale (an example worthy of an undergraduate textbook in Landscape Ecology). For this reason these two general plant communities (grassland and marsh biomes) were treated simultaneously and included in this location in Range Types of North America. The tule marshes were also included under the Meadows chapter of the Grassland biome to facilitate use by students.

In the Arkansas River lowland example employed below differences in salinity and moisture conditions of soil at small (local) spatial scale resulted in a small-scale mosaic of wet (and somewhat saline) tallgrass prairie, mixed prairie, and bulrush or tule marsh in amazingly close proximity to each other.

167. Wet Saline Prairie- Tallgrass prairie comprised of switchgrass, the dominant, and big blusestem, the associate species, surrounded and, in turn, was surrounded by more saline areas dominated by inland saltgrass (Distichlis stricta= D. spicata var. stricta) with associated species ranging from the forb, frogfruit (Lipppia lanceolata) which was the broadleaf species in foreground of this slide, to plains lovegrass (Eragrostis intermedia) to Japanese brome (Bromus japonicus) and, with much less cover, cheatgrass (B. tectorum). Both of these Bromus species are naturalized Eurasian cool-season annuals.

The stand (a consociation) of switchgrass (the bottomland form; a lowland eoctype) seen in background background was shown at local scale in the next photograph. The local stand of tallgrasses (mostly switchgrass) in left foreground was on a more saline microhabitat so that grass growth and development was less and behind that of the same species on less saline soil.

Clarification: It was explained in several captions below that grassland vegetation on the Arkansas River lowland that was described in this section was labeled as wet prairie or wet saline prairie and not marsh. Designation of marsh was applied only to wetlands having their land surface covered with water during most--at least during critical parts--of the plant-growing season, and typically supporting grasslike plants rather than grasses. The tule or bulrush marsh presented later provided an example of marsh vegetation. Such marshes are not tallgrass prairie (they are not grassland at all), but the example referred to was included (below) in this portion of the Tallgrass Prairie chapter because these herbaceous wetlands occur in restricted areas within the tallgrass prairie region.

General floodplain (lowland) of the Salt Fork of Arkansas River. Salt Plains National Wildlife Refuge, Grant County, Oklahoma. June; late vernal aspect. Range plant communities of tallgrass species (mostly big bluestem and switchgrass) was FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM rangeland cover type 710 (Bluestem Prairie), but situated within this range community were communities of short- and/or midgrass species, especially inland saltgrass. These latter grassland communities often covered greater area than surrounding tallgrass vegetation. There was not an SRM inland saltgrass rangeland cover type. Bluestem "Tall-Grass" Series 142.11 of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Saline Subirrigated range site. Central Great Plains- Prairie Tableland Ecoregion, 27d (Woods et al., 2005).

168. Standing together- Stand of tallgrass prairie, of which a bottomland ecotype of switchgrass was dominant and big bluestem was associate to local co-dominant, on higher ground of a wet prairie (floodplain of Arkansas River). This was a "photo-quadrant" of the range vegetation shown in the background of the immediately preceding photograph. Tallgrass stands like this one had developed on the higher level (elevation) land whereas lower ying local relief commonly supported consociations of inland saltgrass. Certain plant species such as plains lovegrass, Illinois bundleflower, and both japanese chess and cheatgrass (two naturalized Eurasian annual grasses) were more common in local ecotones (edges) between with these two distinct range plant communities. This extensive vegetational mosaic existed around freshwater tule (bulrush) marshes that had stands of eastern cottonwood (Populus deltoides) around their perimeters. This spatial arrangement of freshwater marshes and a "patchwork" of tallgrass wet prairie interspersed with midgrasses on the greater floodplain of the Arkansas River rproduced a landscape-scale grassland-marsh complex with interacting ecosystems.

169. Tallgrass and midgrass; mound and swale- Two views of local stands of tallgrass wet prairie (switchgrass and big bluestem, with the former more commonly dominant) on slighted elevated ground (including mima mounds as shown here) interspersed with local, primarily single-species stands (consociations) of inland saltgrass on lower elevation land. Differences in elevation of land (soil surface) was usually just a matter of inches to a few feet even from depth of swales or "micro-valleys" to top of mima mounds, but it was sufficient to account for the profound differences in these local range plant communities. Salt content was undoubtedly a major factor in determining which of these two drastically distinct range plant communities grew on the two forms of local relief (micro-topography). Other edaphic factors were almost assuredly involved also, including greater soil depth on mjma mounds.

The first photograph presented a view of this range at a camera distance that showed the overall vegetational mosaic and "lay of the land" with both mima mound and low-lying intemound spaces. The second photograph was taken at shorter camera distance and showed specifics of the local relief and corresponding range vegetation (eg. switchgrass and big bluestem on the same mima mound and inland saltgrass all around base of teh mima mound). Relatively large areas of bare soil surface were widespread and characteristic of this wet prairie range.

Illinois bundleflower and plains lovegrass were locally abundant at outer edges of the stands of midgrass (mostly inland saltgrass) such as shown growing conspicuously in center and right foreground in the second slide. Western ragweeed (Ambrosia psilostachya) was an associate species on such perimeters (lower right corner of second slide).

Question as to proper designation of wetland: It was not known whether this natural wetland vegetation was more precisely described as wet prairie or as salt marsh. Perhaps the tallgrass (switchgrass and big bluestem) range vegetation was wet prairie and the slightly lower elevation and more saline soil (usually a consociation of inland saltgrass) was salt marsh. This author was reluctant to label any grassland plant community as a marsh instead restricting the designation of marsh only to land inundulated with water (standing water on the land surface) for a good part of the plant-growing season (eg. the tule or bulrush marsh covered below).

170. Salty stand- Wet saline prairie in part of the Salt Fork Arkansas River Valley that was a consociation of inland saltgrass that extended over a large area as part of a vegetational mosaic with tallgrass (switchgrass and big bluestem) prairie that developed on slightly land. A large expanse of this tallgrass plant community was in the far background of this photograph. About the only other plant species in this expansive stand of inland saltgrass was Japanese brome or Japanese chess, and it was present only as scattered individuals.

Successional state of the inland saltgrass community was not known, but this worker suspected that it was climax, or at least the potential natural vegetation for which a climax might not exist if the soil in the swale (the fairly level land situated within or among mima mounds) was not a mature soil. The description of this range site in the soil survey (Soil Conservtion Service, 1985, p. 55) stated that "continued overgrazing and extreme climatic conditions" ultimately results in range deterioration to a degraded community including inland saltgrass, ragweed, and annual grasses. The pattern o f retrogression on this range site (includinga brief description of intermediate successional stages) may well be an accurate visualization of the path of range depletion, but that range site description did not--indeed, by itself, could not--explain simultaneous presence of extensive populations (single-species stands) of inland saltgrass immediately adjacent to the obvious tallgrass climaax of switchgrass and big bluestem.

It appeared to this rangeman that both the tallgrass and the inland saltgrass communities were climax . Any overgrazing (it would have been in the somewhat distant past as this grassland was not being overgrazed and had not been overgrazed in recent years) would have resulted in replacement of the obvious tallgrass climax on mima mounds the same as on intermound spaces. Thus, it seemed highly unlikely that inland saltgrass was other than climax range vegetation (ie. a consociation). The key words in the range site description by agency range conservationists (Soil Conservtion Service, 1985, p. 55) were most likely "extreme climatic conditions". In more precise ecological terms these three words would be read as something like "harsh microclimate" (ie. "climatic conditions" would refer to microhabitat, microsite, or microenvironment) where much of the "extreme" nature of conditions would edaphic and/or topographic (perhaps reflecting drainage, salt accumulation, and related factors). Presence of climax (decreaser) Illinois bundleflower along with "weedy" seral forbs like western wheatgrass in stands of inland saltgrass was further evidence of the climax nature of range plant communities dominated by inland saltgrass. It was illogical to conclude that overgrazing had depleted tallgrasses and resulted in their replacement by inland saltgrass on land of level or flat microtopography while right next to this microland form tallgrass (switchgrass and big bluesetm-dominated) vegetation on mima mounds had not been impacted by grazing, or had recovered from past grazing abuse so much faster. Some other factor(s) had to be more responsible than grazing management.

General floodplain (lowland) of the Salt Fork of Arkansas River. Salt Plains National Wildlife Refuge, Grant County, Oklahoma. June; late vernal aspect. FRES No. 41 (Wet Grasslands Ecosystem). No Kuchler or SRM designation for inland saltgrass. In Brown et al. (1998, p. 45) this would be Saltgrass Series 242.34 (if one was shown which it was not) of Plains Interior Marshland 242.3. Saline Subirrigated range site. Central Great Plains- Prairie Tableland Ecoregion, 27d (Woods et al., 2005).

171. Saltier stand- Grassland landscape in the Great Salt Plains of northcentral Oklahoma. The general range plant community of this area was tallgrass prairie, but highly saline soils resulted in large areas dominated by halophytes, plants adapted to saline habitats, situated within the regional tallgrass prairie climax.

This was an example of how and where concepts of Landscape Ecology conveniently described/explained the spatial organization of these range plant communities.

These two wide-angle photographs constituted a "nested photoplot" or "nested phototransect" in which the second or lower photograph was a smaller area within the overall grassland landscape presented in the first or upper slide. In these two slides, there were large, sprawling stands of inland saltgrass (Distichlis stricta) and Pursh seepweed or horned sea blite (Suaeda depressa= S.calceoliformis) with alkali sacaton (Sporobolus aeroides) forming the outer edges of these stands that formed on highly saline edaphic environments. In terminology of Landscape Ecology, the patches of saline grassland had developed within a matrix of tallgrass prairie). The areas of bare soil were apparently so salty that even these halophytes could not establish.

Alkali sacaton also formed a more-or-less continuous margin or zone around the tallgrass prairie as well as the saline shortgrass vegetation.

This was another example of where the polyclimax theory of vegetation development by seemed more descriptive or explainative than the older monoclimax theory of Clements (1916).

172. Salty standouts- Cover of inland saltgrass and Pursh seepweed or horned sea blite that formed extensive stands or patches of halophytic grassland within the surrounding matrix of tallgrass prairie in the Great Salt Plains of northcentral Oklahoma. This area is the western portion of the Central Lowlands physiographic province or the Pririe-Plains Border (Fenneman, 1931, ps. ). cal

173. On the salty edge- Alkali sacaton formed the outer margin of Four Horsemen (big bluestem and switchgrass were dominant to little bluestem and Indiangrass) tallgrass prairie on the Great Salt Plains in northcentral Oklahoma. Alkali sacaton also formedd a vegetational zone at outer edges of numerous stands or patches of inland saltgrass and horned sea blite, a shortgrass form of range vegetation, as shown in the immediately preceding two-slide set.

There were also widely spaced plants of willowleaf baccharis (Baccharis salicina) scatered in this tallgrass prairie community.

Salt Plains National Wildlife Refuge, Grant County, Oklahoma. Range plant communities of tallgrass species (mostly big bluestem and switchgrass) was FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM rangeland cover type 710 (Bluestem Prairie), but situated within this range community were communities of short- and/or midgrass species, especially inland saltgrass. These latter grassland communities often covered greater area than surrounding tallgrass vegetation. There was not an SRM inland saltgrass rangeland cover type. Bluestem "Tall-Grass" Series 142.11 of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Saline Subirrigated range site. Central Great Plains- Prairie Tableland Ecoregion, 27d (Woods et al., 2005).

174. Still salty- Stand of tallgrass prairie dominated by big bluestem and switchgrass and with willowleaf baccharis forming sporadic woody cover in this climax grassland vegetation. These two slides comprised another "nested photoplot" or "nested photoquadrant" with the second slide being a photographic subplot with the larger "photoplot" of the first slide. Several comparatively short plants of willowleaf baccharis were present in the smaller subplot of the second slide.

These two photgraphs were taken at outer margin of the tallgrass community such that there was also substantial cover and biomass of alkali sacaton which was common along the outermost edges of both this tallgrass prairie and shortgrass vegetation comprised of inland saltgrass and horned sea blite.

175. Fresh water bulrush or tule marsh- General or overall views of a freshwater marsh at edge of the Arkansas River Valley (Salt Fork) existing as a consociation of tule or bulrush known variously as Americn bulrush, saltmarsh bulrush, Olney threesquare, and chairmaker's club-rush, (Scirpus olneyi= S. americanus= Schoenoplectus americanus) and with the twining forb, fogrfruit (Lipppia lanceolata), as associate species. Minor (other than locally) plant species included American water plantain (Alisma subcordatum), southern annual saltmarsh aster (Aster divaricatus= A. subulatus var. ligulatus), and Engelman's spikerush (Eleocharis engelmannii). For all practical purposes there were essentially no other plant species present in this freshwater marsh.

The first of these two slides presented a wider view of the entire range plant community of the tule marsh and associated range vegetation around the perimeter from adjoining plant communities. Range communities around the margins of the marsh wer shown in the second slide. This second photograph showed a local forest of eastern cottonwood that had developed around edges of the marsh. This local forest had been invaded by the naturalized and dreadfully invasive shrub, Russian olive (Elaeagnus angustifolia), that was readily distinguished by its silvery gray-colored leaves.

Edge species: in this immediate locality there were places where sedges (Carex and Cyperus spp.) and rushes (Juncus spp.) grew along perimeters of freshwater tule marshes where this range vegetation contacted wet--often saline--prairie. An example of such contacts and the resulting local-scale ecotonal (transitional) vegetation, including sedges and rushes, was shown at end of this section.

Salt Plains National Wildlife Refuge, Grant County, Oklahoma. June; late vernal aspect. FRES No. 41 (Wet Grasslands Ecosystem). K-42-Tule Mrashes. No SRM designation of a rangeland cover type for tule marshes. In Brown et al. (1998, p. 45) this would be Bulrush Series 242.33 (if one was shown which it was not) of Plains Interior Marshland 242.3. Central Great Plains- Prairie Tableland Ecoregion, 27d (Woods et al., 2005).

176. Stand of chair-makers' rush ( Scirpus americanus)- The local freshwater marsh introduced in the preceding two photographs was a consociation of a major species of bulrush that has a "pasal" of both common and scientific names including Olney threesquare, chairmaker's club-rush, saltmarsh bulrush or Americn bulrush (Scirpus olneyi= S. americanus= Schoenoplectus americanus). Frogfruit was the associate--and about the only other--plant species of this marsh other than incidental (found only sporatically) species which were listed in the immediately preceding caption. It was possible that there were some infrequent plants of hardstem bulrush (Scirpus acutus= Schoenoplectus acutus), but this species could not be positively identified.

Note on taxonomy of Cyperaceae including the bulrushes or tules: Radical changes in nomenclature and general taxonomic organization have been in the Cyperaceae that have created confusion, if not chaos, among all users of scientific names except for the elites who change the names and systematics. To ease the pain and facilitate study of range vegetation both the traditional and revised (ie. revolutionary) binomials were shown for this species.

Detailed views of this wetland range vegetation were presented in the next two slides and caption.

177. Those of a freshwater marsh- Two progressively closer views of range vegetation in a freshwater marsh almost exclusively dominated by chairmakers' club-rush, chair-makers' rush, Olney threesquare, or Americn bulrush. The associate species was frogfruit (mostly visible in the second slide). Other--though only incidental--species included American water plantain, southern annual saltmarsh aster, and Engelman's spikerush. The chairmakers' or American bulrush was in early bloom stage. (An example of inflorescence and stem of this bulrush species was presented below.)

178. Tule marsh and pond (and cows in it)- A vegetational mosaic of tallgrass (switchgrass and big bluestem-dominated) wet prairie, inland saltgrass saline prairie, and freshwater marsh dominated by (a consociation of) American bulrush or chairmakers' rush vegetation on floodplain (lowland) of the Salt Fork of Arkansas River. These three distinct range plant communities were shown and described above (in this section, entitled Continuum of Wet Prairie to Marsh). The tallgrass (switchgrass-big bluestem) wet prairie and the inland saltgrass low (saline) prairie existed in such intiricate and intimate association that they could be viewed as one general grassland (a biome) community that was distinct from the marsh (another, even if small-scale, biome). That these two (if viewed at biome-scale and distinction) or three (if viewed on basis of dominant plant species) range plant communities sometimes developed "cheek by jowl" was shown cleaerly in this photograph. In fact, a natural (not manmade) freshwater pond (with two cows) in the tule marsh was thrown in for good measure.

Range vegetation in foreground was transitional (ecotonal) between tallgrass wet prairie and tule (bulrush) marsh. Major range species in this local ecotone included switchgrass (a bottomland ecotype), Torrey rush (Juncus torreyi), Ehgelmann's spike-rush, caric sedges (Carex spp.), and umbrella sedges (Cyperus spp.). Species of the latter two genera could not be identified in their current vegetative (pre-bloom) phenological stages.

Grant County, Oklahoma. June; late vernal aspect. FRES No. 41 (Wet Grasslands Ecosystem). K-42-Tule Mrashes. No SRM designation of a rangeland cover type for tule marshes. In Brown et al. (1998, p. 45) this would be Bulrush Series 242.33 (if one was shown which it was not) of Plains Interior Marshland 242.3. Central Great Plains- Prairie Tableland Ecoregion, 27d (Woods et al., 2005).

179. Bulrush and cordgrass- Freshwater marsh on margin of salt plains in Central Lowlands physiographic province in northcentral Oklahoma. The dominant range plant was American bulrush (known also as chair-makers' bulrush, chair-makers' rush, and Olney threesquare) with prairie cordgrass (Spartina pectinata) and tapertip flatsedge or tapertip umbrella-sedge (Cyperus acuminatus) as associates. Major forbs were spotted water hemlock (Cicuta maculata) and blue vervain (Verbena hastata). Pink thoroughwort, pink Joe Pye-weed (Eupatorum incarnatum) was a less abundant forb.

This wetland did not typically have an inundated or covered soil surface yearlong, but the soil was saturated most of the year so that this was a marsh, an herbaceous wetland. This freshwater marsh was situated within the overall tallgrass prairie. In the framework of Landscape Ecology, freshwater marshes were patches within the matrix of tallgrass prairie.

Salt Plains National Wildlife Refuge, Grant County, Oklahooma. Late July; peak standing crop (or nearly so). FRES No. 41 (Wet Grasslands Ecosystem). K-42-Tule Mrashes. No SRM designation of a rangeland cover type for tule marshes. In Brown et al. (1998, p. 45) this would be Bulrush Series 242.33 (if one was shown which it was not) of Plains Interior Marshland 242.3. Central Great Plains- Prairie Tableland Ecoregion, 27d (Woods et al., 2005).

180. Botanical players on a wetland stage- Two views of a freshwater marsh in the salt plains area of northcentral Oklahoma showing principal plant species and structure (along with botanical composition). In the first or upper slide American bulrush (background), overall dominant species was shown with tapertip flatsedge (foregound) the associate species.

The second slide featured spotted water hemlock and blue vervain, the two major forbs in the freshwater marsh community.

Salt Plains National Wildlife Refuge, Grant County, Oklahoma. Late July; just about all species at peak standing crop and full bloom. FRES No. 41 (Wet Grasslands Ecosystem). K-42-Tule Mrashes. No SRM designation of a rangeland cover type for tule marshes. In Brown et al. (1998, p. 45) this would be Bulrush Series 242.33 (if one was shown which it was not) of Plains Interior Marshland 242.3. Central Great Plains- Prairie Tableland Ecoregion, 27d (Woods et al., 2005).

A major, multi-named, and confusing bulrush- Dense stand (first slide) and flower cluster near end of sexual shoot (second slide) of an aquatic grasslike plant known variously as American bulrush, chairmaker's rush, common threesquare, Olney's bulrush, and Olney's threesquare (among others). These examples were growing in a freshwater marsh in the Salt Plains area of tallgrass prairie in northcentral Oklahoma. This plant was the overall dominant species of this wetland.

The triangular shape of the shoot of this Scirpus species was obvious in the second slide.

Salt Plains National Wildlife Refuge, Grant County, Oklahoma. Late July; fruit-ripe stage of phenology.

181. Taxonomy and nomenclature: Recent (beginning primarily in 1990s) names and interpretations of phylogenetic relations in the bulrushes or tules is a bloody mess. According to the unpublished Great Salt Plains Refuge plant species list (and referencing back to published species synonyms) these species of Scirpus were on this range:. 1) chairmakers' rush, American bulrush, or three square (S. americanus), 2) salt marsh or alkali bulrush (S. paludosus= S. maritimus),and 3) soft-stem or great bulrush (S. validus). S validus was shown on the refuge list as synonyous with S. tabernaemontani which has usually been shown as synonymous with S. acutus which was generally known as a separate species with common names of hard-stem or, also, great bulrush. Thus there might be as many as four species of bulrush or tule on the range of this refuge. Finally, it was noted that all of these tule or bulrush species which were previously shown as Scirpus had been changed to Schoenoplectus.

These examples of chairmakers' rush, American bulrush, or three square were from Cherry County, Nebraska. Late June (near peak biomass, some shoots at full-boom).

182. "Froggie Went A-Courtin'"- Frog (sometimes, fog)-fruit (Phyla lanceolata) is a member of the vervain family (Verbenaceae) that grows on a remarkably diverse array of habitats. These range environments generally seem to be locally disturbed and/or wet microsites. One seemingly inconsistent habitat was as an associate species on a tule marsh in northcentral Oklahoma at the western edge of the the tallgrass prairie region. Specimens of frog-fruit there were not as advanced as others previously photographed by the author so the latter were "transplanted" here. Either way and on both ranges, frog-fruit fared better than Froggie (or was it "Froggy"?) and Miss Mouse in the timeless ballad.

Ottawa County, Oklahoma. June; full-bloom stage.

183. Spotted deadly- Spotted waterlock (Cicuta maculata) shown as whole herbacous view (first slide) and upper shoots complete with inflorescence (second slide) growing in a freshwater marsh in the Salt Plains of northcentral Oklahoma. This plant was growing in association with American bulrush or chairmaker's rush and blue vervain. Larger shoots of all these wetland plants extended to heights of seven (or, rarely, more) feet.

This wetland range community was a botanically diverse (and interesting) assemblage of plants.

Technical note: The second slide was over-expoded due to lens malfunction or, more accurately, due to improper lens maintenance. Unbeknownst to the photographer, an incompetent servicemen had applied white grease as a lens lubricant years earlier when the lens underwent routine preventive maintenance! Yes,"believe it or not" the servicemen were downright stupid and used tool grease in a camera lens. Wonder is that the ole Nikon/Nikoor macrolense had worked for years and thousands of slides before it eventually could no longer perform properly. Fortunately there are competent servicemen and one of these restored the old Nikkor to its original proper functioning. Taken in conjuction with other photographs of spotted water hemlock, students atill get a composite view of spotted watr hemlock.

Salt Plains National Wildlife Refuge, Grant County, Oklahoma. Late July; full-bloom phenological stage.

184. Poisonous on the plains- Upper shoots with umbels (first slide) and compound leaves (second slide) of spotted water hemlock growing in a freshwater marsh in the Salt Plains of northcentral Oklahoma.

Spotted water hemlock is one of the most toxic of North American range plants. In fact, many toxicologists and plant taxonomists regard it as the single most poisonous native to North America. The poisonous principle is cicutoxin, a naturally occuring alcohol. This is a species of large, rank-growing plants that are generally unpalatable to just about everything, However, numerous deaths--to both livestock and humans--have from ingestion of spotted water hemlock. Hence, "cowbane"is one comon name that reflects this toxicity to livestock.

Human deaths are almost always due to accidental poisoning when the immense roots of this member of the parsely family are mistakenly eaten for "carrots" or "parsnips". When handling plants of this species (as in photography of range plants) all body parts that came in contact with them should be washed thoroughly and as soon as possible.

Salt Plains National Wildlife Refuge, Grant County, Oklahoma. Late July; full-bloom phenological stage.

185. Toxic details- Main stem and branches of spotted water hemlock growing in a freshwater marsh in the Salt Plains of northcentral Oklahoma. As is typical of many species of the Umbelliferae, the carrot or parsely family, spotted water hemlock is a biennial or, less commony, a perennial. Also typical of the umbelliferous species are the large, bulbous roots of this virulently poisonous plant.

Salt Plains National Wildlife Refuge, Grant County, Oklahoma. Late July; peak standing crop phenological stage.

186. No spots on flowers- Umbel (upper slide) and clusters of flowers in umbel (seond slide) of spotted water hemlock growing in a freshwater marsh in the Salt Plains of northcentral Oklahoma.

Salt Plains National Wildlife Refuge, Grant County, Oklahoma. Late July; peak standing crop phenological stage.

187. Non-poisonous with poisonous- Upper shoot of blue vervain (Verbena hastata) accompanied by American bulrush and spotted water hemlock growing in a freshwater marsh in the Salt Plains of northcentral Oklahoma.

This is one of the larger, taller Verbena species. It prefers wet to moist habitats like that of this wetland.

Salt Plains National Wildlife Refuge, Grant County, Oklahoma. Late July; full-bloom phenological stage.

188. Blue-topped- Upper shoots (first slide) and flower cluster or inflorescence (second slide) of blue vervain growing in association with American bulrush or chairmaker's rush and spotted water hemlock. Individuals of this species grow to comparatively large size (heights of seven or more feet on this marsh).

Forage value of blue vervain to livestock is essentially zero. It is, however, a good bee plant. In Range Management one never assumes that a species of low palatability to livestock is a weed or competitor to "valuable species". It may be that species like blue vervain provide certain features, factors, or habitat conditions that are beneficial or even essential to more palatable range species.

Salt Plains National Wildlife Reguge, Grant County,Oklahoma. Late July; full-bloom phenological stage.

189. Wet prairie- First bottoms floodplain tallgrass prairie dominated by prairie cordgrass (also called sloughgrass) with eastern gamagrass and bottomland switchgrass as two associate species. The conspicuous forbs are foxglove beard-tongue (Penstemon digitalis) which is nearing end of its flowering period and swamp milkweed (Asclepias incarnata) which is at late pre-bloom stage. Almost no grasslike plants such as Carex, Juncus, or Scirpus species were common.

The heavy clay soil of this bottomland retains so much moisture that prairie crawfish have dug numerous shafts and thrown up short earthen chimneys throughout the ground of what is one of the largest and most pristine examples remaining of this rare type of tallgrass prairie. So far this priceless representative of the once vast North American prairie has been protected by it's use as an unbelievably productive hay meadow.

Cherokee Prairie sub-province of the Central Lowland physiographic province. This is a part of the general Osage Section of the overall Central Lowland province.

Cherokee County, Kansas. Vernal aspect, June. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 601(Bluestem Prairie), more generally, or SRM 710 (Bluestem Prairie). Central Irregular Plains- Cherokee Plains Ecoregion, 40d (Chapman et al., 2001).

Organization note: examples of foxglove or smooth beardtongue were shown below with other members ot the Scrophuloraceae, the snapdragon family, including other Penstemon species..

190. Swamp milkweed (Asclepias incarnata)- Swamp milkweed on go-back ground (an old field) on wet prairie in the western Ozark (springfield) Plateau. Ottawa County, Oklahoma. August.

191. Swampy pollinators- An assortment of pollinating insects, including both those of orders Diptera (first slide) and Lepidopetra (second and third slides) visiting the swamp milkweed plants introduced in the preceding slide.

Photographic note: certain butterfly species, including monarch (Danaus plexippus) have their wings in almost constant motion while feeding. Thus photographs taken at slow speed (like 1/15 second as done here) to get depth of field will often show the blurred moving wings like those in the monarch in third slide.

Ottawa County, Oklahoma. July.

192. Wet prairie- First bottoms floodplain tallgrass prairie dominated by eastern gamagrass with bottomland switchgrass the general associate species (locally the dominant) with prairie cordgrass the third major grass species. This was a separate hay meadow adjoining the one shown in the last photograph of wet prairie, but here prairie cordgrass came in third behind the other two bottomland prairie species.

This was another example of one of the most beautiful bottomland tallgrass prairies the author was ever blessed to enjoy.

Cherokee Prairie sub-province of the Osage Section of the Central Lowland physiographic province. Cherokee County, Kansas. Vernal aspect, June. FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 601 (Bluestem Prairie), more generally, or SRM 710 (Bluestem Prairie).

193. Wet spot- A seasonally (typically spring and early summer) wet prairie in the western Ozark (Springfield) Plateau populated almost exclusively by two caric sedges, fox caric sedge (Carex vulpina) and low caric sedge (C. brevior). Both species were at peak standing crop and with ripening achenes.

This one local habitat was all that remained of a tallgrass prairie that decades earlier had been converted to crop land and then planted to tall fescue (Festuca arundinacea) and bermudagrass (Cynodon dactyledon). This parcel of wet prairie remained because it was too wet to farm.

Ottawa county, Oklahoma. Early June; late vernal aspect. Not a good identifier set for this range vegetation.

Nebraska Sandhills (and related postclimax tallgrass types)

Sand and sandy soils (also gravel for that matter) have a higher proportion of macropores (noncapillary pores) so that there is less water runoff and more water that infiltrates this rooting material for plants. Conversely,there is less water retained in the coarse or heavy textured soils as compared to fine (light)-textured soils such as clay; however, , more of the retained water in coarse textures (eg. sand) is available for plant absorption. Also many plants that are adapted to coarse-textured soils have a high proportion of their roots near the land surface so that they absorb this temporarily shallow water before it evaporates or percolates below their root zones. Net result is that the native vegetation on sand, sand dunes, and high sand-content soils is typically more mesophytic than vegetation on adjacent or neighboring soils comprised of less coarse particles (those that have finner soil texture).

This was explained in Weaver and Clements (1938, p. 203-204) by the Clementsian concept of chresard. Chresard is that part of total soil water, the whole content of water in soil (= holard), that is available for plant absorption or use by the plant part. The echard (to withold) is that portion of water still present (held or retained) in soil at the permanent wilting point (ie. water that is unavailable for plant use; water that adheres to tightly to soil particles that plants cannot absorb it). Holard minus chresard equals echard (or any other arrangement of these portions ssuch as holard minus echard equals chresarde or chresard plus echard equals holard). In the Clementsian monoclimax model natural vegetation of sandy environments if postclimax "In Nebraska the relation [transition from true to mixed prairie] is further disturbed by the extensive sand-hill region, in which the high chresard favors a postclimax of tall grasses far beyond their proper climate [ie. beyond the climatic or regional climax]" (Weaver and Clements, 1938, p. 524). "In the vast sand-hill area of Nebraska, the tall-grass postclimax attains its best development, which is assumed to reflect the climate when the prairies were occupied by the bluestems and their associates some millions of years ago" (Weaver and Clements, 1938, p. 86). This also included shrubs such as New Jersey tea, sand sagebrush (Artemisia filifolia), and soapweed yucca.

The sandhill region of Nebraska and similar smaller areas in other parts of the Great Plains including some in Colorado, Kansas, Oklahoma, Texas, and New Mexico support more mesic (more moisture-requiring) range vegetation than that of less porous soils in the same region or area. In the vernacular of Tansley's polyclimax model or Whittaker's climax pattern interpretation postclimax vegetation would be edaphic and/or topographic climax vegetation (edapho-topographic climaxes). Same ecological (successional or climax) outcome.

Tallgrass prairie range vegetation in the Nebraska Sandhills (and some similar range communities in Colorado, Oklahoma, New Mexico, Texas, etc.) is in the Mixed Prairie Region though, of course, it is not mixed prairie. As such, postclimax or edaphic-topographic climax tallgrass prairie was covered in this chapter. This included such range types as sand sagebrush- and shinnery oak (Quercus havardii)-tallgrass savannas as well as the classic Nebraska Sandhills tallgrass prairie. Range types of mixed prairie, including some in the Nebraska Sandhills (not all Sandhills grassland is tallgrass prairie), were (logically) dealt with in the chapter, Mixed Prairie.

Organization note: The Nebraska Sandhills as well as related tallgrass prairie subtypes or forms of the Southern Great Plains (= High Plains, Staked Plains, Llano Estacado) were treated separately in the subsequent chapter, Tallgrass Prairie (Interior)- II. A few photographs of the famed Nebraska Sandhills were retained in this chapter to make for a smoother transition to that next chapter that was devoted to tallgrass prairie vegetation of the semiarid zone.

194. Farmed Sand Hills of Nebraska ("God's Own Cow Country")- This western-most extension of tallgrass prairie is typically a community of the Four Horsemen species. Region-wide little bluestem, State Grass of Nebraska, is clearly dominant in an abiotic environment more characteristic of tall and true prairies than the mixed prairie that would be expected in this semiarid precipitation zone. The Sand Hills range vegetation type is the product of soils of deep sand, often accompanied by shallow surface acquifers. In classic Clementsian view this tallgrass prairie is postclimax to both true and mixed prairie. This is a Choppy Sands range site with sand bluestem (Andropogon hallii), sand lovegrass (Eragrostis trichodes), and prairie sandreed (Calamagrostis longifolia) the dominant species. Cherry County Nebraska. Hiemal aspect, October. FRES No. 39 (Prairie Ecosystem). K-67 (Nebraska Sand Hills Prairie). SRM 602 (Bluestem- Prairie Sandreed). Nebraska Sand Hills- Sand Hills Ecoregion, 44a (Chapman et al., 2001).

195. Sands range site of the Nebraska Sand Hills- Same species as above with more mid and short grasses like blue and hairy grama (Bouteloua gracilis,B. hirsuta) and Junegrass. Cherry County, Nebraska. Hiemal aspect, October. FRES No. 39 (Prairie Ecosystem). K-67 (Nebraska Sand Hills Prairie). SRM 602 (Bluestem- Prairie Sandreed). Nebraska Sand Hills- Sand Hills Ecoregion, 44a (Chapman et al., 2001).

196. Sandhills Marsh- Wetland range site.Mostly grass-like plants such as rushes (Juncus spp.), cattail (Typha angustifolia, T. latifolia), bur-reed (Sparganiumeurycarpum ) locallywith sedges (Carex spp.) throughout.Note lodge of muskrat (Ondrata zibethica). Cherry County Nebraska. As part of tallgrass prairie this is in FRES No. 39, but as a marsh it could as logically be placed in the tremendously varied FRES No. 41 (Wet Grassland Ecosystem). Either a hydric variant of K-67 (Nebraska Sand Hills Prairie) or K- 42 (Tule Marshes). Wetlands variant of SRM 602 (Bluestem- Prairie Sandreed). Nebraska Sand Hills- Lake Area Ecoregion, 44d (Chapman et al., 2001).

Final reminder: complete treatment of Nebraska Sandhills and other tallgrass prairie range plant communities of the semiarid zone are in the chapter, Tallgrass Prairie (Interior)-II.

Directions to related range types- Several other examples of tallgrass prairie or tallgrass savanna wetlands were included within Range Types of North America. It was (always will be) arbitrary as to where these (and other) range cover types should be situated within a publication. Inclusion at this juncture within this Tallgrass Prairie chapter was logical, but this chapter was already "overloaded". Likewise some of these wetlands though comprised of tallgrass species (among others) had developed as "islands" within the semiarid precipitation zone and not in the Tallgrass Prairie Region. The most obvious of these were some, including marshes, within the Nebraska Sandhills. These were therefore covered in the chapter, Tallgrass Prairie (Interior)-II.

A wet (essentially subirrigated throughout much of the year) prairie or prairie savanna dominated by bottomland switchgrass (Panicum virgatum) and Maximillian sunflower (Helianthus maximillianii) with narrowleaf cattail (Typha domingensis), heath aster (Aster ericoides), and pink boneset (Eupatorium incarnatum) as associate herbs and with black willow (Salix nigra) as a woody component had developed in an ecotone of Grand (Fort Worth) Prairie and West Cross Timbers. Treatment of this wetland range vegetation was included in a section entitled, "Savanna Wetlands" in the chapter, Meadows and Related Marshes, under the heading,Grasslands.

Also in the Meadows and Related Marshes chapter and under the section entitled, Savanna Wetlands, was a wetland savanna dominated by bottomland switchgrass, American bulrush or chairmaker's rush square (Scirpus americanus= S. olneyi), and hairy (hairy seed) rose mallow (Hibiscus lasiocarpos) with eastern cottonwood (Populus deltoides) and black willow (Salix nigra) in the Cherokee Prairie of northeast Oklahoma plus a savanna of eastern cotton and peachleaf willow (Salix amygdaloides) with great or soft-stem bulrush (Scirpus validus), tapertip flat (umbrella) sedge (Cyperus accumulatus), narrowleaf cattail (Typha domingensis) in the Smoky Hills of central Kansas.

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