Tallgrass Prairie (Interior)-I

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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).

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). 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).

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." Tless 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. 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). Limy Upland range site. Flint Hills Ecoregion, 28a (Chapman et al., 2001).

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. 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). Clay Upland range site. Flint Hills Ecoregion, 28a (Chapman et al., 2001).

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. 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).

15. 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. 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).

16. Sward of upland tallgrass prairie- Two "photo-quadrants" of the south slope big bluestem-switchgrass Flint Hills range introduced in the preceding slide. 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. 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).

17. 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! 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). Loamy Upland range site. Flint Hills Ecoregion, 28a (Chapman et al., 2001).

18. 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. 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). Clay Upland range site. Flint Hills Ecoregion, 28a (Chapman et al., 2001).

19. 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).

20. 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. 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).

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.

21. 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).

22. 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).

23. 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.

24. 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, May. Vernal aspect, May. 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).

25. 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. Vernal aspect, May. 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).

26. 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).

27. 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).

28. 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). 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).

29. 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).

30. 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).

31. 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).

32. 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).

33. 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).

34. 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).

35. 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).

36. 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).

37. 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).

38. 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.

39. 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).

40. 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).

41. 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).

42. 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.

43. 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).

44. 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).

45. 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).

46. 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).

47. 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).

48. 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).

49. 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).

50. 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).

51. 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).

52. 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).

53. 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).

54. 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).

55. 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.

56. 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. Diamond Grove Prairie, Newton County, Missouri, July.

57. 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).

58. 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).

59. 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).

60. 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).

61 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.

62. 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.

63. 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).

64. 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).

65. 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).

66. 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). Bluestem "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).

67. 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). Bluestem "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).

68. 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. 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). Bluestem "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).

69. 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 Bluestem "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).

70. 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).

71. 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- Juniper-Oak Savanna. Texas. Variant of SRM 717 (Little Bluestem-Indiangrass-Texas Wintergrass). Cross Timbers- Limestone Cut Plain Ecoregion 29e (Griffith et al., 2004).

Texas Blackland Prairie

72. 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).

73. 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).

74. 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).

75. 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).

76. 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).

77. 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.

78. 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).

79. 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. 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).

80. 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. 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).

81. 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. 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).

82. 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.

83. 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.

84. 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).

85. 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).

86. 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).

87. 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. 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).

88. Ozark Prairie-Another view (in another year) of the same hay meadow described in the two 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).

89. 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. 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).

90. 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).

91. 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. 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).

92. 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). 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).

93. 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.

94. 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. Great 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).

95. 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. General floodplain (lowland) of the Salt Fork of Arkansas River.Great 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). 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).

96. 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). General floodplain (lowland) of the Salt Fork of Arkansas River. Great 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).

97. 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. Great 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).

98. 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. Great 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).

99. 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. 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).

100. 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.) Great 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).

101. 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).

102. 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).

103. "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.

104. 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..

105. 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.

106. 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.

107. 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).

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.

Dominant, Prominent, Major, and Otherwise Interesting Range Plants

of the Tallgrass Prairie

Grasses of Tallgrass Prairie

108. Texas bluegrass (Poa arachnifera) in the Texas Western Cross Timbers- Throughout the southern two-thirds  (or more) of the North American tallgrass, true, mixed, and shortgrass prairies and plains warm-season grasses are far more  pre-dominant and important than cool-season species on basis of species number and biomass production. This is more the case the farther south grassland vegetation extends because numbers of panicoid and eragrostoid species increase while festucoid grasses decrease with southward progression (conversely the proportion of festucoid species increases with northward progression in the continental grasslands). Usually,  however, there are some warm-season grasses in the north and some cool-season grass species in southern grasslands. Texas bluegrass is one of the these southern cool-season grasses. It is one of the more conspicuous and locally dominant festucoid grasses in the southern latitude grasslands. This dioecious rhizomatous species sometimes forms exclusive colonies at microsite scale especially on deep sand sites as seen here in the Cross Timbers of northcentral Texas. These colonies can be bisexual or largely monosexual suggesting that like buffalograss they are monoclonal (consisting of one genotype or genetic individual as a clonal organism). Two things are certain about Texas bluegrass: 1) it is a species that is highly palatable to grazing animals and 2) it is perceived by human eyes to be an extremely attractive plant. In regards the second point, many native plant fanciers rank Texas bluegrass among their favorite prairie species which gives this grass value for natural landscaping. The rancher and landscaper find common cause in this unique grass.

109. Texas bluegrass (Poa arachnifera)- A climax cool season species of the tallgrass prairies and Cross Timbers of central Texas. Prairie hay meadow, Erath County, Texas, April.

110. Girls' git-together- Colony of female sexual shoots of Texas bluegrass. This was a local group of clonal units of the same female plant (one genetic individual) growing in the West Cross Timbers of northcentral Texas.Erath County, Texas, Late April; peak bloom stage..

111. Soparno duet- Two sexual shoots of female Texas bluegrass in the colony introduced in the immediately preceding photograph. Can there be any doubt that this is, simply, one of the beautiful native grasses of southern portions of the Tallgrass Prairie Region? The author has always tried to encourage the wise-use conservation, restoration, establishment, and enjoyment of range plants, especially the most valuable or readily enjoyed native species. This includes such ancillary uses as prairie landscaping and planting of natives as ornamentals. There are few, if any, native prairie grasses in the areas where Texas bluegrass naturally grows that fit the bill any better than this cool-season, dioecious grass. In addition to such obvious uses as forage for livestock and wildlife, Texas bluegrass makes a beautiful lawn species such as the examples shown here. If ya'll are so fortunate as to have in your lawn take note of the obvious: DO NOT MOW IT UNTIL IT GOES DORMANT. Why would any rational person opt to shred off such lovely panicles? It would be the equivalent of a woman with beautiful flowing locks shaving her head. Erath County, Texas, Late Aprill; peak bloom stage.

112. Inflorescence of female Texas bluegrass- The female panicle of Texas bluegrass is larger than  the male, but both are quite attractive. Texas bluegrass shares the common diagnostic feature of the florets of Poa species: the "cobwebby"lemma of the pistillate florets due to presence of cottony like hairs on the callus of the lemma. The specific epithet, arachnifera, refers to arachnid or spider. Erath County, Texas. April.

113. Showy Texas ladies- Female spikelets of Texas bluegrass. A view that preented the beauty of Texas bluegrass in bloom. The pistillate spikelets are bigger and more showy. Girls are supposed to be showier (sweeter, too). Erath County, Texas. Mid-April.

114. Male plant of Texas bluegrass- Flowering of male Texas bluegrass just before anthesis. Vernal aspect, April. Young County, Texas.

115. Junegrass- Prairie hay meadow, Burkhart Prairie, Newton County, Missouri. June.

116. Inflorescences of Junegrass- Tallgrass prairie hay meadow. Newton County, Missouri. June

117. June-bloomers- Junegrass (Koleria cristata= K pyramidata= K. macrantha) growing with porcupinegrass (Stipa spartea) and broomsedge bluestem (Andropogon virginicus) on tallgrass prairie in western Ozark Plateau. Occurrence of Junegrass with porcupinegrass on this tallgrass prairie indicated the ecological affilitation of tallgrass prairie with true prairie. Junegrass is festucoid species in of the oat tribe (Aveneae) and has a semi-contracted or narrow panicle. Junegrass is a much more important species in the true and mixed prairies than in tallgrass prairie. Clements (1920, ps. 121-124) interpreted the true prairie as the Stipa-Koleria Association with Junegrass one of five species forming consociations (though this was less common for Junegrass). It was Junegrass' widespread distribution and association with most other common grass species that distinguished K. cristata as a dominant of true prairie which, even at the time Clements (1920) described it, had been largely eliminated by farming. Clements (1920, ps 131-134) and Weaver and Clements (1938, ps.520-521) regarded the bluestem-Indiangrass (Andropogon species as then interpreted)-dominated grasslands eastward of true prairie as subclimax prairie, the Adropogon associes. Today, Clements' subclimax is described as the tallgrass prairie (Weaver and Clements, 1938, 520-521). Junegrass has also long been regarded as a dominant of the mixed prairie (Clements, 1920, ps. 135-138; Weaver and Clements, 1938, p. 523) in which it forms consociations. Clements (1920, p. 138) regarded Junegrass as about mid-range in the array of dominants with regard to water requirements. In essence, Junegrass is one of the most widely distributed native grasses theoughout North America, occurring from Canada's Northwest Territorities to Quebec and south and east to Alabama, but it almost never abundant other than as local consociations. Newton County, Missouri. Early June; peak bloom stage.

118. Narrow pyramids- This three-slide sequence presented at consecutively closer camera range examples of the narrow or semi-constricted panicle with two-flowered spikelets of Junegrass at anthesis. These panicles were on the same tallgrass prairie as in the preceding slide. This meadow had been used for production of prairie hay for close to a hundred years. The arthropod atop the first panicle was a female American dog tick (Dermacentor variabilis) patiently waiting for her victim. Vegetation in humid and subhumid regions is often literally crawling with ticks such as this specimen throughout much of the summer. The photographer left this one hungry, but she should have not been completely disappointed as she got this publicity shot. Newton County, Missouri. Early June; peak bloom stage.

119. Done for the year- Fully mature and sensecent plant of Junegrass with panicles full of ripe grain. Junegrass is a perennial bunchgrass, a cespitose (tufted or bunched) species. The first slide presented five separate plants while the second photograph showed a single individual to emphasize the bunched habit of this member of the oat tribe. All of these plants were growing on a relict of tallgrass prairie in the western Ozark Plateau that was within the Prairie Peninsula. An oak-hickory forest was just behind the small glade-like opening of prairie that was prime habitat for these two and a half foot tall plants. These photographs were taken five days--and on a drier microhabitat--later than those in the two immediately preceding slide sets. Newton County, Missouri. Early June; grain-ripe and early dormancy stage.

120. Ripe and ready for shedding- Narrow panicles of Junegrass with two-floret spikelets at grain-ripe/shatter stage. The group of panicles shown in the first photograph were on the plant presented in the last preceding photograph immediately above. The second photograph was a close-up of a portion of one of these semi-contracted panicle. Small tallgrass prairie situated within an oak-hickory forest in the Prairie Peninsula (western Springfield Plateau). Newton County, Missouri. Early June; grain-ripe/shatter phenological stage.

121. Another cool member of the oat tribe- Prairie wedgegrass or wedgescale(Sphenopolis obtustata) on a tallgrass prairie in the Prairie Peninsula of the western Ozark Plateau. This approximately 60 acre prairie had been used as a hay meadow for nearly a century or, perhaps even longer as its history of haying was lost with the ghosts of haymakers past. Big bluestem was the dominant of this meadow vegetation. The cool-season prairie wedgescale could survive with the larger, more competitive, more water use-efficient members of the bluestem tribe (Andropogoneae) by completing its life cycle before the tallgrass species got much more than started on the rapid- growth phase of their annual cycles. This striking and conspicuous plant is an uncommon--though a widespread--festucoid species. prairie wedgegrass is most at home on the virgin sod of central prairies, but it grows from Maine across to British Columbia and from the Sonoran and Chihuhuan Deserts across to northern peninsular Florida. Newton County, Missouri. Early June; late anthesis.

122. Cool-season panicles on the prairie- Sexual shoots with panicles (first slide) and a single (individual) panicle of prairie wedgescale or wedgegrass growing on a tallgrass prairie in the Prairie Peninsula. Grasses are by-and-large determinate bloomers meaning that flowering begins with the uppermost and outermost spikelets (in contrast to lowermost and innermost flowers in indeterminate species). In panicles seen here flowering had advanced to the lowermost spikelets with the most distal (uppermost or "highest") spikelets having completed flowering and ferttilization and progressed to soft-dough grain stage. Newton County, Missouri. Early June; late anthesis.

123. Wedged spikelets- Details of a branch of the panicle (first photograph) and opened florets on this branch (second photograph) of prairie wedgescale or prairie wedgegrass. This was a lower branch off of one panicle such as the one presented immediately above. The open (blooming) florets of spikelets on this panicle branch were the upper- or outermost ones indicating that this particular unit of the inflorescence had just begun anthesis. This was a closer-in look at determinate flowering. Newton County, Missouri. Early June; early anthesis.

124. Sexually explicit- Details of florets in spikelets on a branch of a panicle of prairie wedgescale. Anthers and stigmas were exerted (ie. "letting it all hang out" in the vernacular of The Hombres). Newton County, Missouri. Early June; early anthesis.

There are a number of wildryes (Elymus spp.) that are important (even locally dominant) cool-season perennial grasses on tallgrass prairie. Three of the more common and widespread species in the heart of "tallgrass country" and adjoining hardwood forests and savannahs (including the famed Prairie Peninsula) were presented below. Elymus species ae in the wheat or barley tribe (Tritaceae or Horedae).

125. Stand of Canada wildrye (Elymus canadensis)- Large, dense population of Canada or nodding wildrye growing on a floodplain of a branch of the Bosque River in the Western Cross Timbers-Grand Prairie vegetational area of northcentral Texas. Many shoots in this beautiful, luxuriant stand were bent over from rush of recent flood waters. "Pure" (single-species) stands of this size are quite rare because this species does not usually form populations or colonies over large areas as do some species on tallgrass parairie (eg. big bluestem, little bluestem, prairie cordgrass, purpletop). Erath County, May; peak standing crop.

126. Nodding along an ehemeral stream- Several plants of Canada or nodding wildrye with characteristic features and growing in its "prime habitat", moist to wet soil in a fairly cool, shaded locale. These plants were part of the large stand of this species that was shown immediately above growing on a floodplain of the Bosque River. This is the typical habit of this species with the large spike inflorescences "nodding" (drooping or bending over at peduncle, the juncture of inflorescence and rest of shoot). Erath County, May; peak standing crop.

127. Starting to nod over- Examples of spikes of nodding or Canada wildrye that were beginning to bend down (or over) at stage of anthesis. These flower clusters will continue to bend farther downward as the sexual shoots of these plants get ever drier near end of their annual cycle and as spikelets in these spikes get heavier with growing, ripening grin. Erath County, May; anthesis.

128. Virginia wildrye (Elymus virginicus)- A few plants of Virginia wildrye had grown to large size on a protected microsite of a tallgrass hay meadow on the Springfield Plateau of the Ozark Mountains. These plants were growing on the backside of a motte of persimmon where hay-making equipment could not enter. Late haying (August or September) over a span of nearly half a century had greatly reduced decreaser tallgrass species on this meadow, but dormancy (usually) or low-growing, early shoots (sometimes) at late summer harvest permitted plant of this cool-season species to thrive amid fierce grassland competition. Ottawa County, Oklahoma. June (early summer); late anthesis to milk stage of grain.

129. Spikes of Virginia wildrye- Sexual shoots of the Virginia wildrye plants that "paraded" in the immediately preceding photograph. Awns of Virginia wildrye are conspicuously shorter than those of other Elymus species typically found on tallgrass prairie and Prairie Peninsula ranges. Compare relative lenghts of these awns to those of the species introduced before and after E. virginicus. Ottawa County, Oklahoma. June (early summer); late anthesis to milk stage of grain.

Some more examples- Shown immediately below were more photographs of shoots and spikes (whole-inflorescence shots) of several individuals of Virginia wildrye growing in an oak-hickory savanna in the Ozark Plateau just east of the Cherokee Prairie of the Central Lowlands physiographic province. Ottawa County, Oklahoma. July.

130. Study of an understorey grass- Three-slide sequence showing sexual shoots of Virginia wildrye (Elymus virginicus) in forest understorey in western Ozark Plateau. Peak standing crop with grain in mid-dough phenological stage. Ottawa County, Oklahoma. July.

131. Spikes in the shade- Examples of spikes of Virginia wildrye in understorey of western Ozark Plateau forest at mid-dough stage of phenology. Ottawa County, Oklahoma. July.

Another wildrye- Presented next was silky wildrye (Elymus villosus), another locally abundant cool-season species on tallgrass prairie. Specimens shown in this short section were growing on the same range as the examples of Virginia wildrye presented immediately above. These two very similar Elymus species can be distinguished based on curved vs. not curved glumes (Kucera, 1961, p.73; Kucera, 1998, p. 111), but they can be differentiated even more quickly on the range based on complete emergence of the spike from the boot in E. villosus in contrast to completely exerted (and generally larger) spikes in E. virginicus. (It is remarkable how often specialists overlook obvious, stark differences and instead "split hairs" on minutae.)

132. Silky wildrye (Elymus villosus)- When discussing the central grasslands of North America most students of the range typically refer to Canada or nodding and Virginia wildryes as the major Elymus species. A lesser known or recognized wildrye growing over much of the tallgrass prairie, Prairie Peninsula, and open understorey oak- hickory forest is E. villosus, silky wildrye. With the longer-awns and larger spike of nodding wildrye and the vertical or "straight" spike of Virginia wildrye silky wildrye looks very much like a hybrid of the two more commonly recognized prairie species. Silky wildrye is, however, a species "in its own right" and one that is often much more common than the two better known Elymus species. These plants and those presented immediately below were growing in a part of the Prairie Peninsula at the western perimeter of the Ozark Plateau. Ottawa County, Oklahoma. June (late spring); peak anthesis.

133. The beauty of reproduction- To the agricultural producer nothing is as exciting as the promish of reproduction, be it a developing, pregnant female or blooming "ears" of grain. Infloresecences of silky wildrye at peak anthesis are a vivid example of the sentiment expressed by Senator Ingalls of Kansas when he wrote that the "homely hue" of grass was "more enchanting than the lily or the rose". Ottawa County, Oklahoma. June (late spring); peak anthesis.

134. Another native, cool-season, prairie grass- Winter bentgrass (Agrostis hyemalis) on moist to wet local habitat on tallgrass prairie in western Ozark Plateau. This is a minor species other than in local mesic microsites. It was included here to give an idea as to the great species diversity of tallgrass prairie. Even in southern areas native, winter, grasses grow alongside warm-season, perennial, panicoid grasses. Winter bentgrass has generally been regarded as a short-lived perennial (Gould, 1975, p. 140). It grows on both interior and coastal tallgrass prairie. Ottawa County, Oklahoma. Late May, maturing grain to early grain-ripe phenological stages.

135. Dainty panicles on tallgrass prairie- Panicles and spikelets of winter bentgrass. These flowers were on some of the same plants shown in the immediately preceding slide. Ottawa County, Oklahoma. Late May, early grain-ripe stage..

136.. Two examples of little bluestem colonies- Little bluestem, prairie beardgrass, broom beardgrass, or common beardgrass (Andropogon scoparius= Schizachyrium scoparium) was probably the single most important range grass across pre-Columbian North America. This was likely true based on geographical range of the species, the total quantity of animal feed it produced (say, measured as Animal Unit Months), and the land area (acreage) on which this species was the dominant or co-dominant plant species. Hitchcock and Chase (1951, p. 754) showed little bluestem as occurring in every state of the Union except Nevada, California, Oregon and Washington. Little bluestem is found in the three Canadian Prairie Provinces (Looman and Best,1987, ps. 96-98) and as far eastward in Canada as Ontario. It occurrs in the Chihuhuna Desert, but not in the Sonoran Desert. With loss of most of the original tallgrass, true, and mixed prairies (and the more favorable sites therein) to the plow much of the original cover of little bluestem was destroyed. Less of the virgin range dominated by blue grama (Bouteloua gracilis) was plowed so a shortgrass species that was substantially less widespread and lower-yielding has now likely surpassed the bluestem and other tallgrass prairie species in importance as a range plant. If the contemporary interpretation of "little bluestem" is accepted such that this is a complex of taxa that were formerly interpreted as being distinct species the enlarged S. scoparium may still be the most important range plant in North America based on the criteria specified above. Examples of bluestems or beardgrasses formerly treated as distinct species by agrostologists like Hitchcock and Chase (1951) that were reinterpreted as subspecies or varieties and "lumped" in the enlarged, "umbrella" S. scoparium by more recent taxonomists like Gould (1975) and Allen (1992) included pinehill bluestem (Andropogon divergens= S. scoparium var. divergens), seacoast bluestem (A. littoralis= S. scoparium var. littoralis), New Mexico little bluestem (A. neomexicanus= S. scoparium var. neomexicanum). Little bluestem often forms naturally occurring single species stands, the Clementsian consociation, over large areas and is a co-dominant (or at least a major species) on many range sites. Dense populations of this least mesic of the Four Horsemen of the Prairies species often occur as vast natural "fields". Such stands are extremely productive of biomass and serve as examples of why defoliation of herbage whether by grazing animals or fire is so important to mainteance of grassland. The heavy yields of biomass and high density of tillers in these two populations will result in excessive accumulations of dead plant material unless reduced by grazing and/or fire. The build-up of excess mulch will effective exclude light from basal portions of the little bulestem plants the following spring and delay "green-up" and retard plant growth and herbage production during the ensuing growing season. Both stands of little bluestem shown in these photographs were in the Western Cross Timbers and Prairies vegetational area of northcentral Texas (Erath County) and at peak standing crop. The stand in the first slide was on a Bottomland range site in late fall (the latter part of October). The stand in the second slide was on a Rolling Prairie range site in anthesis during early fall (late September).

137. Little bluestem plants- The cespitose habit of little bluestem was illustrated by these plants growing on a Rolling Prairie range site in a portion of the Grand Prairie in the Western Cross Timbers and Prairie land resource area of Texas. Unlike big bluestem, Indiangrass, and switchgrass (the other three of the Four Horsemen tallgrass species) little bluestem is not rhizomatous. All secondary shoots are intravaginal (vertical shoots designated as tillers or often, among farmers and stockmen, "stools"). As such little bluestem is never a sod-forming grass although individual plants (genetically distinct individuals) frequently grow so close together as to form a prairie sod or dense sward.

138. Single plant of little bluestem- Little bluestem is the one common dominant grass of the tallgrass prairie that is strictly cespitose (ie. a bunchgrass or tussock grass). Older individual plants form large tussocks like this one which had a basal cover of over two and a half feet feet. This growth is in effect asexual reproduction by increase in numbers of tillers (intravaginal, hence upright, shoots). Little bluestem also flowers prolifically (shown in slides below), but this sexual reproduction is generally not as efficient as asexual reproduction. This is especially the case in established swards. Most of the tillers of this plant had advanced phenologically to become flowering shoots. Portion of Grand Prairie on Tarleton State University Hunewell Ranch, Erath County, Texas. September.

139. Flowering shoots of little bluestem- Both anthers and filaments of stamen as well as stigma were visible on the sexually reproductive tillers of little bluestem presented in these two slides. Tarleton State University Hunewell Ranch, Erath Cpounty, Texas.. September.

140. Anthesis in little bluestem- This close-up view of staminate and pistillate organs in little bluestem was presented as another example of flower structure in the Gramineae. Tarleton State University Hunewell Ranch, Erath County, Texas. September.

141. Inflorescences of little bluestem- Sexually reproductive shoots of little bluestem at seed-ripe stage. Inflorescences of the bluestem or sorghum tribe (Andropogoneae) have typically been interpreted as racemes. Chase (1964, p. 82) described the arrangement of the Andropogon flower cluster as having racemes "borne on numerous slender leafy branches arising in the axils of leaves on the main culm or branches, the whole forming a compound inflorescence". Other workers like Highnight et al. (1988, p. 8) labeled this inflorescence type as a spicate raceme. The rachis is jointed and falls apart upon maturity resulting in the shattering of grains. Tarleton State University Hunewell Ranch, Erath County, Texas. September.

142. Raceme of little bluestem- This spicate raceme had advanced to the seed-ripe stage just before the next phenological stage of seed-shatter. The spikelets are paired: one perfect and sessile, the other sterile and pedicellate, with two florets per fertile spikelet. These two florets (in the fertile spikelet) are as follows: one floret is perfect and terminal and the other (the lower) floret is present only as a sterile lemma. The paired spikelets fall entire and together, the sterile pediceled spikelet attached to the fertile sessile spikelet, from the jointed rachis of the compound spicate raceme. The individual branches of this compound raceme are often defined as a rame, an inflorescence branch which bears some pediceled and some sessile spikelets. Two entire rames and the basal part of a third rame were displayed in this slide. Tarleton State University Hunewell Ranch, Erath County, Texas. September.

143. Basal portion of a shoot of little bluestem- It was explained above that little bluestem is strictly a cespitose species (a bunchgrass) the tillers (upright, intravaginal shoots) of which form a tussock habit. In spite of the rank or relatively coarse and large size of little bluestem the individual tillers are rather easily broken off at the ground level (ie. at or just above the root crown) of little bluestem plants. This was shown in these two photographs. In other words, even though this species evolved under heavy grazing and is well-adapted to defoliation even it can be damaged-- remarkable easy-- by imporper grazing, especially overuse (overstocking). In time, prolongued overuse leads to overgrazing and a change in species composition of the range plant community. Increasers and invaders are much less easily damaged than the tallgrasse species including little bluestem. On a portion of the Grand Prairie on Hunewell Ranch, Erath County, Texas. September.

144. Colony of big bluestem- Big bluestem is the dominant tallgrass species across much of the tallgrass (= bluestem) prairies that once extended from western Ohio and the Canadian Prairie Provinces to the central Dakotas and south through the Blackland Prairie and coastal prairie region of Texas, specially on soils having calcareous parent materials like limestone and dolomite. Big bluestem produces more sexually reproductive shoots under spring burning regimes as shown hereon the Konza Prairie (Riley County, Kansas, July).

145. Color namesake of big bluestem- This is an example of spring coloration for which bluestem (big bluestem in particular) was named. Technically the blue pigmentation is at tips of leaves not stems but blueleaf just does not have the poetic ring of names so characteristic of the frontier and rural folk in general. Big bluestem is generally the number one dominant tallgrass species (first overall among the Four Horsemen of the Prairies in the Tallgrass Prairie Region). Obviously dominance, abundance, and related ecological characteristics are range site-specific with different climax grass species being dominant on different habitats. Commonness, relative abundance, palatability, adaptation to fire, etc. of big bluestem are responsible for this species often being regarded as "king of the tallgrass prairies". Ottawa County, Oklahoma. Estival aspect, June.

146. Basis of the king-grass species- Characteristic basal shoots of big bluestem on tallgrass prairie in western Ozark Plateau. Pubescence and semi-flattened culms that are typical of big bluestem could be seen in detail in these three photographs. Hitchcock and Chase (1950, p. 757) described the pubescence on basal leaves of big bluestem as being occasionally densely villous (long, soft hairs). Hitchcock and Chase (1950, ps. 749-751) divided the Andropogon genus into three sections based on numbers and other features of the spikelet-bearing branches which they termed racemes. Big bluestem was included in the Arthrolophis section which included such highly unpalatable and invader species as broomsedge bluestem (A. virginicus), splitbeard bluestem (A. ternarius), and bushy bluestem (A. glomeratus). Contemporary treatment in Barkworth et al. (2003, ps. 652-664) limited Andropogon to the Arthrolophis section of Hitchcock and Chase (1950, ps. 750-751) and divided this restricted Andropogon genus into two sections: 1) Andropogon which included only big bluestem and sand bluestem (A. hallii) and 2) Leptopogon with all the other Andropogon species of the traditional Arthrolophis section. Andropogon and Leptopogon sections were distinguished/segregated based on features of pedicellate spikelets (in effect, well-developed versus vestigial or wanting, in the respective sections). Big bluestem is what has been called variously a "culmless", "short-shoot", or "short-culmed" grass. This king of the tallgrass prairie--overall the dominant, single most important, and most palatable of the Four Horsemen of the Prairies--is a tallgrass species that does not elevate its "growing point" (apical meristem) until relatively late in the warm growing season. Big bluestem blooms about as soon as the sexual shoots have elongated. This is in contrast to Indiangrass and switchgrass ("culmed", "long-shoot", or "long-culmed" grasses) which elongate their shoots early in the growing season and, therefore, much earlier than onset of flowering. Ottawa County, Oklahoma. Late June; young vegetative state with apical meristems still low.

147. Inflorescence of big bluestem with the characteristic three rames (inflorescence branches bearing some pedicellate and some sessile spikelets) from which arrangement is derived another common name of turkey track. Ottawa County, County Oklahoma, August.

148. Rame of big bluestem- The anthers and stigmata were shown fully exerted on these spikelets of big bluestem at anthesis. Newton County, Missouri. September.

149. Autumn coloration of big bluestem- This colony of big bluestem at peak standing crop and seed-shatter stage illustrated the phenomenon of seasonal colors of the prairie. The grasslands-- most notably, tallgrass prairie-- of North America undergo color changes with progression of the seasons and phenological stages of the plants. This is the equivalent of seasonal color change in the deciduous forests that were often (sometimes still are) conterminous and intermixed with tallgrass-dominated grasslands and savannas. These are the seasonal aspects that have been noted periodically in this segment of the publication. Autumnal aspect of tallgrass prairie was pronounced in this stand of big bluestem that was growing near the western extent of it's geographical range in central Texas. The ecotype shown here was scarcely four feet in height (four foot woven wire in immediate background) which was no taller than stands of little bluestem adjacent to this colony. (The far background was woody invasion of live oak, post oak, and mesquite on overgrazed tallgrass Grand Prairie range). Mills County, Texas. October.

150. Sexually reproductive shoots of big bluestem- Grassses are regarded as reproducing by both sexual and asexual means. The latter is essentially multiplication of secondary shoots (tillers, rhizomes, or stolons) by vegetative growth (hence asexual reproduction is also known as vegetative reproduction). Perennial grasses typically reproduce more prolifically (and more effeciently/effectively) by vegetative reproduction than by sexual (grain production) means. This is most pronounced in many of the prairie grasses native to North America. Flowering and fruit (caryopsis) production in these species is essentially a "luxury" action or physiological response. Going back to an early classic sutdy by Branson (1953) various authors have placed different grasses species into various groups based on certain characteristics. One of the more common of these groupings was a scheme based on location and timing of elongation of the apical meristem in combination with proportion of shoots that became sexually reproductive (Heady and Child, 1994. p.23). These authors placed big bluestem in the group described as having "infertile apices numerous and in or near the soil". The relatively high proportion of big bluestem shoots that were developed infloresences and produced grains in this colony resulted from a summer that was considerably cooler and slightly wetter than was typical. Mills County, Texas. October.

151. Inflorescences of big bluestem- Rames of big bluestem at the seed-ripe stage and in autumn coloration. Individual spikelets along the rachises were conspicuous in these unusually fertile and heavy yielding shoots. Mills County, Texas. October.

152. King and Queen of the Prairies- Big bluestem (right) and little bluestem (left) on a portion of Grand Prairie in the West Cross Timbers and Prairies vegetational area of northcentral Texas. This was at the western edge (the least subhumid or least mesic part) of the species range for big bluestem in this land resoruce area (ie. habitat was marginal for big bluestem). Consequently big bluestem, King of the Prairies, had already matured and was at the seed-ripe (and fast approaching seed-shatter) stage while little bluestem, Queen of the Prairies, was still in the soft dough phenological stage.At this geographic location and on this range site (Laomy Prairie) the range environment was "prime habitat" for little bluestem, regional dominant of tallgrass prairie in central Texas, and this species was slightly later (slower) in it's annual cycle and lagged behind big bluestem, the general dominant tallgrass species across or "averaged over" the entire bluestem-Indiangrass prairie (such Kuchler units as K-66 and rangeland cover types as SRM 710, both designated Bluestem Prairie). Astute observers will have noted that the ecotype of big bluestem growing on habitat marginal for this species produced shoots no larger (taller) than those of little bluestem. This phenomenon was not the general or typical condition relative to respective sizes of these species on tallgrass prairie. The usual size dimorphism was embodied by common names of the two species. Mills County, Texas.

153.. Indiangrass- The State Grass of Oklahoma is the major co-dominant with the bluestems, especially big bluestem, of the tallgrass prairie range type. Indiangrass is more abundant and the dominant grass on soils derived from sandstone parent material and is thus relatively more common south of the zone where big bluestem is the major dominant. Indiangrass is typically the tallgrass dominant over much of the Osage Questas and particularly the Chautauqua Hills portions of the Central Lowlands physiographic province south of the Flint Hills section. As shown here both species are herbaceous dominants in the forests and savannas of the Ozark Plateau section. Newton County, Missouri, October.

154. Colony of Indiangrass- These five to six feet-plus shoots of flowering Indiangrass at peak standing crop in a fencerow show the remarkable drought-tolerance of native tallgrass species. This biomass yield was at summer’s end in one of the most severe droughts in Texas history. It grew during the fourth year of a protracted drought, the last two growing seasons of which were drier than any two of the great drought of the 1950s. Shoots in this colony remained green throughout the entire duration of each growing season in the prolonged drought of the 1990s. Grand Prairie vegetation. Erath County, Texas. October.

155. Inflorescence of Indiangrass- This panicle (in anthesis) illustrates the resemblance of the flowering shoot of a dominant prairie grass to the arrow atop the head of an Indian brave and thus origin of the name Indiangrass. (By the way, the author finds it obvious that the first letter in this common name should properly and always be capitalized given that Indian is a proper noun. Incidentally, Indian— and not that horribly offensive, sickeningly affected, and incorrect invention “Native American”— is the correct, precise name for American aborigines.)

156. Graminaceous headress- Panicle and flag leaf (first photograph) and spikelets on branches (second photograph) of yellow Indiangrass. The more specific common name of yellow Indiangrass is often aplied to Sorgastrum nutans in contrast to slender Indiangrass (S. elliottii) and lopsided Indiangrass (S. secundum). The latter occurs only sporatically from North Carolina to Louisiana and Arkansas except in Florida where it is widespread. Branches with spikelets arising from the cental axis is the pattern or arrangement of main floral units is a panicle which strictly speaking is the only form of compound inflorescence in the Gramineae. The flag leaf is the leaf immediately subtending (below) the inflorescence in grasses. Hunewell Ranch, Erath County, Texas. October; peak-bloom stage (and almost simultaneous anthesis in all spikelets).

157. Beauty in flowers of Oklahoma State Grass- Close-up "shots" of Indiangrass spikelets in anthesis.As chance or Divine Intervention would have it, this and the preceeding pair of photographs showed almost all spikelets in anthesis at once. This flowering sequence is unusual in most grass species because grasses are determinate bloomers (flowering proceeds from upper and outer spikelets downward and inward). Indiangrass is one of the most prolific grain-producers of the native tallgrasses and, as such, is one of the better native species for reseeding of tallgrass, true, and mixed prairie ranges (Phillips Petroleum Company, 1963, p. 62; Leithead, 1971, p. 157; Tyrl et al., 2008, p. 163). In fact, Indiangrass, big bluestem, and switchgrass have, in recent years, been "rediscovered" as warm-season species for use in tame (agronomic) pastures where extensive management is possible and desirable. Hunewell Ranch, Erath County, Texas. October; peak-bloom stage (and almost simultaneous anthesis in all spikelets).

Details of two of the Four Horsemen grasses- Close-up photographs of big bluestem and Indiangrass were presented immediately below. Both of these tallgrass species are in tribe Andropogoneae and have the tribal characters of: 1) paired spikelets on a jointed rachis with one spikelet perfect and sessile and the other spikelet being sterile and pedicellate, 2) the fertile spikelet, in turn, has one perfect floret that is terminal and a sterile (empty) lemma below the perfect floret, 3) the fertile spikelet (with its paired florets; one perfect, one sterile) is shed along with the rachis joints, and while 4) the sterile and pedicellate spikelet remains attached (Chase, 1964, p. 99). In other words, there are paired spikelets with the fertile one of these spikelets having paired florets with the uppermost one of these paired florets bing fertile (perfect). Both florets (one fertile, one sterile) of the perfect and sessile spikelet are shed as a unit (the spikelet) while the sterile spikelet remains on its pedicel which stays attached to the inflorescence. Examples were growing side-by-side on an oak-hickory-tallgrass savanna on the western edge of the Ozark (Springfield) Plateau. Details of these two members of the Andropogoneae were presented together for consistency in describing fetures of this panicoid tribe.

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158. Next genetic generation of big bluestem- Details of sexual propagules of big bluestem at scale of spikelets. Close-up vies of rames of big bluestem showing individual spikelets of this champion species of the tallgrass prairie and tallgrass-eastern hardwood savanna. Rame is the traditional term applied to branches of the panicle of Andropogoneae with there being both pedicellate and sessile spikelets on such branches. (Pohl, 1968, p. 242). (Similarly,the adjectives ramose, meaning branching or having many branches, and rameal, in reference to oroccurring on a branch, are used in standard taxonomic talk.) Paired spikelets--both perfect and sessile and sterile and pedicellate--were visible upon close examination of rames in both of these photographs. The appendages that appear as (look like) little branches with missing units are the sterile, pedicellate spikelets and not an empty space where a spikelet or floret was attached. However, to add to confusion (as if it was needed), some pedicels were without sterile lemmas (ie. some pedicellate spikelets were present only as pedicels). There were also some whole pedicellate spikelets that did have their one sterile lemma. Generally, intact fertile (perfect) and sessile spikelets were still attached. Some of all floral units were present so that the complete arrangement of paired spikelets and paired florets of perfect spikelets were still attached. Phenological stage was grain-ripe not grain-shatter. Stoney Point Savanna, Ottawa County, Oklahoma. January; grain-ripe stage.

159. Next genetic generation of Indiangrass- Details of sexual propagules of Indiangrass at scale of spikelets. Sections of panicle branches with mature, grain-filled, fertile spikelets. Paired spikelets--both perfect and sessile and sterile and pedicellate--were visible upon close observation of these two photographs. Sterile, pedicellate spikelets were visible as pedicels with pilose pubescence. Fertile, sessile spikelets were much more obvious in these taken-on-the-range shots. Stoney Point Savanna, Ottawa County, Oklahoma. January; grain-ripe stage.

160.  Silver bluestem (Andropogon saccharoides= Bothriochloa saccharoides= B. saccharoides var. torreyana= B. laguroides subsp. torreyana) at anthesis - This midgrass is typically an increaser on most mixed prairie range sites and an invader on tallgrass prairie range sites, but it is a decreaser on some of the range sites of the arid semidesert grassland or shallower sites in the western Edwards Plateau adjoining the Chihuhuan Desert. Nomenclature of this species seems to be a never-ending source of debate, revision, followed by debate and further revision (ie. another name), none of which adds anything useful or practical to management of this or associated range plants. Erath County, Texas. July.

161. Silver bluestem at seed ripe stage- Characteristic autumn coloration of this prairie midgrass. Note that most of the shoots are sexually reproductive. Erath County, Texas. July.

162. Partly in and out of the boot- The four apices of silver bluestem shoots in these two photographs presented four degrees of emergence of the inflorescence (panicle) from the boot. Panicle atop the left shoot in second photograph was fully emerged or exerted. Details of fully expressed panicles were shown in the next set of two slides. West Cross Timbers near end of a summer of severe drought. Erath County, Texas. September, emergence of inflorescence from boot.

163. Silvery spectacle- No, not necessarily spectacular but a conspicuous spectacle nonetheless was this appearance of showy inflorescences of silver bluestem in the Western Cross Timbers. Terminology applied to inflorescences of Andropogon, Bothriochloa, Schizachyrium, and Dichanthium species has about as many interpretations as does taxonomy of these taxa. Contemorary description of this inflorescence type is panicle with primary racemose branches (Gould 1975, p. 591; Hignight et al., 1988, ps. 8, 23; Hatch and Pluhar, 1993, p.43). These specimens were growing in the West Cross Timbers near end of a long, hot sumer of severe drought and following two mowings. This species is survivor if given half a chance (come to think of, even if not given half a chance). Erath County, Texas.September, fully emerged inflorescence, at 1) pre-anthesis stage (first photograph) and 2) early dough stage (second photograph).

164. Broomsedge bluestem (Andropogon virginicus)- Although broomsedge is closely related to big bluetem (Hitchcock and Chase [1951] placed them in the same section of Andropogon) the former has nothing of the ecological status or forage value of the latter. In fact, broomsedge is an invader throughout the tallgrass prairie (if not everywhere it grows) and it is one of the least palatable of all native perennial grasses throughout its biological range. Individual broomsedge plants form large conspicuous bunches and it is a strictly cespitose species whose shoots are all intravaginated (ie. tillers), but it reproduces very effectively both vegetatively and through abundant seed production. Ottawa County, Oklahoma. Seed-shatter stage; autumnal aspect, early December. Ottawa County, Oklahoma.

165. Tillers of broomsedge bluestem at seed-shatter stage- Spikelets in the specialized inflorescence (a spicate raceme) of broomsedge. Ottawa County, Oklahoma. Early December.

166. Colony of broomsedge bluestem- Broomsedge appears to be a classic example of a ruderal species, a plant associated with man-made disturbances such as waste places or minimally managed land (ruderals are usually viewed as weeds). More specifically, broomsedge was interpreted as a dominant competitive-ruderal that exerts phytotoxic effects (Grimes, 1979, p. 144). Students will get an instructive lesson by studying Grimes' "primary strategies", in this instance of the "low stress-high disturbance strategy" (Grimes, 1979, ps. 7, 39-45). This population of broomsedge was growing in a 40-year old planting of "Kentucky 31" tall fescue. Fescue is a cool-season bunchgrass so it and the cespitose, warm-season broomsedge co-exist (often as co-dominants) on extensively managed permanent pastures of the introduced tall fescue and on old-fields where tall fescue became established as a volunteer crop. This is the Ozark Fescue-Broomsedge Cover Type category of Tall Fescue (SRM 804), an example of which was shown here. In portions of winter through early summer tall fescue will appear as the dominant until it goes into summer dormancy at which time the broomsedge bunches appear as dominants-- at least as an aspect dominants as is also the situation for tall fescue in it's growing season. Broomsedge often forms immense consociations like the one shown here on abandoned farmland and cut-over forests from the tallgrass prairie and oak-hickory uplands of the Ozarks to the Piedmont along the Atlanic Ocean. Broomsedge frequently exists as a midseral stage for prolonged periods on old-fields ("go-back-land") that were depleted by soil erosion and overcropping. Broomsedge is one of the most unpalatable species of native grasses in the tallgrass prairie and oak-hickory-tallgrass savanna. The resemblance (under superficial and careless exmination) of broomsedge to big and little bluestem and it's presence as an invading associate on deteriorated tallgrass prairie was a combination duped many an unsuspecting stockman. At one time many of the large steer operators leased-- and occasionally even bought-- "go-back land" dominated by broomsedge thinking it high-quality bluestem pasture. Jawhawker and Okie landowners had a nice laugh (all the way to the bank) at the expense of steermen (many from Texas). It did not take two such experiences before cattle-rasisers learned that plant identification was not just a hobby for botanists at the local "cow college". Ottawa County, Oklahoma. Autumnal aspect, early December.

167. Basal part of autmunal-hibernal broomsedge shoots- One key part of the specific dominant competitive-ruderal "strategy" (Grimes, 1979, ps. 39-51) of broomsedge bluestem is maintnenace and/or production of green shoots at ground level during what is mostly (= "more-or-less") it's dormant season. Panicoid grasses like the Andropogon species are quite obviously warm-season plants that typically flower and set grain in late summer to fall. The photographs presented here illustrated that broomsedge bluestem reaches seed-ripe and seed-shatter stages in autumn on into early winter. Closer insepection showed that broomsedge maintained (or grew new) live shoots throughout much of the supposed dormant phase of the annual cycle of this perennial grass. This pattern differs from other bluestems such as big and little bluestems and panicgrasses like switchgrass whose shoots die back at or soon after the first heavy fall frosts. In fact, shoots of these decreaser panicoid grasses often enter dormancy before frost. Persistence of live shoots of warm-season species into and throughout much of winter theoretically allows photosynthesis to be extended over a longer period. Location ot this green tissue at the base of the plant where it is largely covered by dead herbage certainly offers it some protection from cold temperatures and freeze damage by infrequent snows, but such coverage also limits photosynthesis. Perhaps the main survival or competitive advantage provided by this growth and pattern of resource allocation is to give this ruderal and ecological invader a "headstart" over the native dominant bluestems, Indiangrass, and panicgrasses (decreasers). The specific function(s) of hibernally green tissue in A. virginicus would be a good research project, and one with practical applications that could help reduce cover and density of broomsedge in both introduced permanent pastures like tall fescue and native tallgrass prairies. Ottawa County, Oklahoma. Early December.

168. Splitbeard bulestem (A. ternarius)- This is another native Andropogon species (in the same section as big bluestem and broomsedge) that is an ecological invader. Splitbeard bluestem does not form immense colonies and occur as a dominant species over such an extensive geographic area as broomsedge, but it is an indicator species on depleted range and reflects past or present mismanagement like overgrazing. This specimen was growing in a 40-year old stand of "Kentucky 31" tall fescue. Ottawa County, Oklahoma. Seed-shatter stage, early December.

169. Tiller of splitbeard bluestem at seed-shatter stage- The annual phenological cycle of splitbeard bluestem coincides with that of broomsedge bluestem. Ottawa County, Oklahoma. Early December.

170. Rames of splitbeard bluestem- Spikelets on two branches of the spicate raceme of A. ternarius. Ottawa County, Oklahoma. Early December.

171. Inflorescence of eastern gamagrass at full anthesis with separate staminate and pistillate spikelets- This resemblance to the tassle of corn or maize (Zea mays)—both species are in tribe Maydeae or Tripsaceae—is origin of the other common name of corngrass. Ottawa County, Oklahoma, June.

172. Staminate florets of eastern gamagrass- Tridens Prairie, Lamar County Texas, June.

173. Pedicellate florets of eastern gamagrass- Tridens Prairie, Lamar County Texas, June.

174. Switchgrass- Specimen of a bottomland ecotype of Panicum virgatum along the banks of the Bosque River.in northcentral Texas Switchgrass occurs as both a bottomland form, as featured herein, as well as an upland form (upland ecotypes). This is the most mesic of the four dominant tallgrass species (Four Horsemen of the Prairies). Switchgrass characteristically grows in lower habitats than the other three dominants, but upland ecotypes hold their own on upland slopes. As of this writing switchgrass is the only one of the four magnificant dominants of the tallgrass prairie that has not been designated a State Grass. Some state proud of its prairie heritage needs to pick it up. Erath County, Texas. September, peak standing crop and immediate pre-bloom stage..

175. Switchgrass- An old field reseeded to Cave In Spring accession of switchgrass released out of Missouri by the Soil Conservation Service. Native warm season, perennial tallgrasses like switchgrass, big bluestem, and Indiangrass have been “discovered” by forage agronomists to be dependable, drought-tolerant, and palatable pasture and hay species which require judicious grazing management but not the expensive intensive inputs of fertilizer, irrigation water, etc. Of course much of this “natural advantage” would be lost if these “wild” species  were  to be domesticated. Newton County, Missouri.

176. Panicles confront the blue- Panicles of a big specimen of bottomland switchgrass at seed-shatter stage stood in beautiful exposure against the cobalt-blue sky of a Texas "blue norther". A "norther" in traditional lingo refers to massive cold fronts that originate as Arctic air masses and move southward often following the Rocky Mountain chain and dispersing over the Great Plains eastward across the prairies of the Central Lowlands physiographic province. Initially the sky of a 'norther" is gray, often a dark charcoal gray or bluish-black, as it meets the warmer, moister air moving northward out of the Gulf of Mexico. After a day or so of battling air masses the colder, drier, northern air prevails over its southern cousin and under dominion of this clear Arctic air the sky turns to azure blue. The "yucky" air so typical of Texas skies in the contemporary atmosphere filled with pollution from the likes of a "zillion" automobile exhausts is "scrubbed" clean as crisp cold air moves out the haze. Such are Kodachrome moments, and when switchgrass stands tall against the air of a "Texas norther" the Lone Star sutterbug captures the meeting of great grass and grand sky. Tillers of this particular switchgrass plant exceeded eight feet in height and were adorned with panicles comprising two feet of that length. Many, perhaps most, caryopses from these compound inflorescences had already shed. Perhaps new plants (genotypes) will join this mighty sexual specimen. Erath County, Texas. Late November, peak standing crop at grain-shatter, late maturity stage of phenology.

177. Big clump- Basal shoots (tillers) of a large specimen of the bottomland form of switchgrass. Switchgrass has relatively long scaly rhizomes, but it forms large bunches or tufts comprised of many robust tillers. Plants of bottomland switchgrass are much larger in contrast to the upland switchgrass form so that the cespitose habit (bunchgrass morphology) of this species is more pronounced, especially under ideal growing conditions. This plant was approaching pre-dormancy stage (note many senescing leaves) in late autumn following an unusually wet period from late summer through end of warm growing season. It showed the immense size of which this species is capable of attaining. Erath County, Texas. Late November, peak standing crop at grain-shatter, late maturity stage of phenology.

178. Big switch- Large panicle on the plant of bottomland switchgrass featured immediately above. It is not uncommon for this expanded panicle to reach lengths exceeding two feet and a half feet and to be nearly a foot across at the base. There are many spikelets along the numerous branches of this inflorescence. Like most of the tallgrass species switchgrass reproduces primarily by asexual means, but it certainly keeps its reproductive options open. his fine crop of florets was produced in an extremely wet late summer and an equally impreseive crop of tiny grain resulted from this profuse blooming. Erath County, Texas. Late September, full-flower stage.

179. Little flowers- Spikelets of the bottomland form of switchgrass. Two views of numerous spikelets in full flower. These floral units were taken off of the large specimen described above. Both anthers and stigmas were exerted, but stigmas were more obvious. Erath County, Texas. Late September, full-flower stage.

180. Stand of beaked panicgrass- Panicum anceps is often a locally dominant species in the tallgrass prairie and, especially, the tallgrass savanna and Cross Timbers. It is a palatable and productive species and often classed as a decreaser depending on range site. This colony grew on a small natural opening (a glade) in the western Ozark Highlands. Newton County, Missouri. September.

181. Up closer- Part of a local stand of beaked panicgrass on mesic to wset tallgrass prairie showing overall features of shoots at boot stage. Western Ozark Plateau (Springfield Plateau section). Ottawa County, Oklahoma. August; boot (immediate panicle-emergence) stage of phenology.

182. Sexual shoots with inflorescences of beaked panicgrass- The flowering shoots of P. anceps on a tallgrass prairie glade in the Ozark Plateau (first slide) and on a wet prairie (second slide). Newton County, Missouri. September; Ottawa County, Oklahoma, August (first and second photograph, respectively) .

183. Beaked spikelets- Arrangement and details of spikelets on beaked panicgrass growing on a prairie in the western Ozark Plateau. Ottawa County, Oklahoma. July: anthesis and pre-anthesis stages.

184. Vine mesquite- Dense stand of vine mesquite showing characteristic sward of this valuable mid-grass. This panicgrass is adapted to a diversity of habitats ranging from understory species in the Texas Cross Timbers through to the rock-strewn shallow ranges of the Edwards Plateau and the floodplain and bosques of semidesert grasslands of the Southwest. Vine mesquite is usually a decreaser and fairly productive of palatable and reasonably nutritious forage when green but when mature it cures with lower nutritive value than associated shortgrasses like buffalograss or blue and black grama. On a lowland site of Windthorst soil, Hunewell Ranch, Tarleton State University, Erath County, Texas, June.

185. Spikelets of vine mesquite- Hunewell Ranch, Erath County, Texas, June.

186. Paniced again- Redtop panicgrass (Panicum agrostoides= P. rigidulum ) growing in a local depression on virgin tallgrass prairie. Redtop panicgrass is one of the lesser-known Panicum species of tallgrass paririe and adjoining savannas (eg. relative to switchgrass, beaked panicgrass, vine mesquite). Even the common witchgrass (P. capillare), an annual species, is better-known (even to most seasoned rangemen) than redtop panicgrass. This is undoubtedly a reflection of the rather restricted habitats (local, low-lying, wet environments) on which this species can survive. On microsites where redtop panicgrass grows this bunchgrass forms small-size consociations with such nearly complete foliar cover that the stand produces a sod-like sward as seen in these two slides. Redtop panic can grow to the approximate heights of upland switchgrass growing on similar habitats. These two species as well as beaked panicgrass all were well-represented on this meadow used for production of prairie hay. This meadow had been improperly hayed (mowed in late summer) for years and had been returned to proper management so as to be in recovery (secondary succession), but management was irrelevant to this local stand of redtop panicgrass that was growing on a swale which ponded water and was to wet to hay. This land had been mowed for hay two weeks prior to time of these photographs. Wet ground permitted the photographing of this by-passed parcel. Ottawa County, Oklahoma. Mid-July, peak standing crop and full-bloom stages.

187. Ya mean this is a bunchgrass?- Redtop panicgrass is a cespitose species (all shoots are tillers), but it commonly grows as such dense colonies in its restricted mesic microhabitats that it appears--at first flush anyway--to be a sod-forming (= stoloniferous and/or rhizomatous) species. The second of these two photographs showed the bunchgrass habit or growth form of this panicoid grass. Ottawa County, Oklahoma. Mid-July, peak standing crop and full-bloom stages.

188. Growing out of the boot- Emergence of immature panicle of redtop panicgrass from the flag leaf, the leaf subtending the infloresecence. Grass inflorescences are pushed out through the sheath of the enveloping last leaf of the shoot as the culm grows (internodes elongate) grow up through it. Ottawa County, Oklahoma. Mid-July, early bloom (maturing panicle) stage of phenology.

189. Top of redtop panicgrass- Entire panicle (first slide) and upper one-third of panicle (second slide) of Panicum agrostoides= P. rigidulum. The branches (secondary shoots) of the panicle of redtop panicgrass occur in whorls around the rachis (sometimes off of just one "side" of the central axis). Ottawa County, Oklahoma. Mid-July, full-bloom (anthesis) stage of phenology.

190. Panicle and spikelets of redtop panicgrass- Upper portion of a mature panicle with nearly ripe grain (first photograph) and of an immature panicle and branches from another maturing panicle at pre-anthesis stage (second photograph). Basis of the common name adjective, "redtop", was apparent in the second slide. By the way, this panicle and these branches were on a rock of chert "growing" on the hay meadow. Chert is a sedimentary rock made up of quartz of the fine granular type in the microcrystalline group (Pough, 1960, p. 231). Chert is the most common of the fine-grained siliceous rocks which when freshly broken has a shiny appearance 470). This stone had been broken and weathered for a number of years in the humid zone of the Ozark Plateau. It resided for decades in a fence row where someone had tossed it to protect sickle-bar mower knives and sections from its hard surface and "just the right size" for causing mower trouble.Chert and related limestones are the main parent material for soils of tallgrass prairie in this region. Ottawa County, Oklahoma. Mid-July.

Rosette panicgrasses- Panicum is the largest genus of the grass family, at least as clasically traditionally treated. In a comprehensive review of the Gramineae, Clayton and Renvoize (1986) generalized that Panicum species have relatively uniform features in spikelet morphology, but otherwise have considerable variability in general morphology (eg. of culms ), anatomy, and physiologiy.Perhaps most fundamental in this context is that as conventionally interpreted Panicum includes both C3 and C4 species; that is, there are. some Panicum species with the Calvin-Benson C3 photosynthesis pathway whereas other Panicum species have the Hatch-Slack C4 pathway of photosynthesis. These species are usually regarded as cool-season and warm-season species, respectively. Hitchcock and Chase (1951, p. 626-636), admittedly consummate "lumpers" at the genus level, sectioned the classic Panicum genus (based on spikelet morphology) into three subgenera based on shoot characteristics. Each of these subgenera was subdivided into untitled sections or subunits. Dichanthelium was one subgenus of Hitchcock and Chase (1951, ps. 627-633). Dichanthelium was divided into 17 sections. In most contemporary treatments of the Gramineae the subgenera of Hitchcock and Chase (1951) were elevated to seperate genera. Dichanthelium--as subgenus or genus--is morphologically characterized by winter rosettes of short, broad leaf blades (though not all Dichanthelium species have this morphological charaacteristic) and secondary branching of the main shoot. These taxa have C3 photosynthesis and lack Kranz anatomy. They are cespitose though some have rhizomes. The main or principal shoot usually terminates in a primary panicle most spikelets of which are chasmogamous (chasmogamous is the adjective that refers chasmogamy, the condition in whith anthers and stigmas are exposed outside of their own flower so that there is potential for cross-pollination). Off of the main shoot there are typically several branches (secondary shoots) which have secondary panicles at least some spikelets of which are cleistogamous, the condition of natural self-pollination (Barker et al., 2003, p. 406). Members of Dichanthelium typically have vernal and autumnal phases (shoot forms), but vernal panicles set "little, if any" fruit. Instead, most grain is borne in the cleistogamous spikelets of the lateral, autumnal panicles (Gould, 1975, p. 478). It could be hypothesized that this cleistogamous flowering (autogamy) is responsible for the many different morphological forms (subspecies, varieties, or whatever) of the different Dichanthelium. Notwithstanding the pronounced differences between Dichanthelium and Eupanicum (the main subgenus in Hitchcock and Chase [1951, ps. 627, 633-636] that does not have rosettes and shoot branching, and does have fertile spikelets throughout) or, in contemporary systematics, Panicum it is by no means clear that these are seperate genera. Even the distinction between cool-season versus warm-season is arbitrary from standpoint of flowering as vernal phase (shoots) of Dichanthelium members actually flower in summer not spring as is the flowering season for typical cool-season species like festucoid grasses. The individual reader can reach his own conclusions. More importantly, it makes no difference from a practical standpoint other than scientific name (and related confusion). There are rosette panicgrasses--Dichanthelium or Panicum--and they are distinct from those related taxa that do not form basal rosettes and branches off of the shoot. There is one thing that all who have worked with these plants can agree upon: they are an identifiction nightmare come true. Plus, they do provide palatable forage and certainly add biological diversity or--and no pun intended--variety to tallgrass prairie range and hay meadow. As such, one of the major species of rosette panics was included below.

Note on organization: several rosette panicgrasses (Dichanthelium taxa) were included in the chapter, Tallgrass Savanna. Most of these examples grew within two to five miles of the example species shown immediately below. In fact some or, even, all of these species might well have grown nearer to each other in this area but the author simply did not find such situation. As found, the species treated below was growing on tallgrass prairie whereas those under Tallgrass Savanna were found growing in a black oak-mixed hickory-tallgrass savannah.

191. Example of the rosette panicgrasses- Local colony of fascicled panicgrass (Panicum lanuginosum var fasciculatum= Dichanthelium acuminatum subsp. fasciculatum) This is one of the principle rosette panicgrass species of tallgrass prairie though less common and widespread than some of the others. It is, however, one of the most distinctive bearing, as it does, one of the most pronounced fascicles in the Dichanthelium group. Ottawa County, Oklahoma. Late July, mid- to near maturity of grain (see below).

192. Vernal fascicles-Apices of the main shoots (not branches) of fascicled panicgrass bearing primary panicles. Actually these leafy units of the shoot are both panicles and clusters or bunches of leaves which by convertion are termed fascicles. Most, if not all, of the Dichanthelium taxa bear such fascicles. Some of the most distinctive and pronounced fascicles are on this species, Panicum lanuginosum var fasciculatum. Ottawa County, Oklahoma. Late July, mid- to near maturity of grain.

193. Cute clusters-Details of some of the fascicles on shoots of Panicum lanuginosum var fasciculatum shown above. Fasicles are clusters or close-bound bunches of major organs particularily as applied to grass shoots with special reference to culms, leaves, or branches of inflorescences (Gould, 1975, p. 632). For example, needles of pines (Pinus spp.) are borne in clusters that are fascicles. Note differences in maturity of spikelets on one shoot. Ottawa County, Oklahoma. Late July, mid- to near-maturity of grain.

194. Fall witchgrass (Leptoloma cognatum= L. cognatum var. cognatum= Digitaria cognata)- Two photographs from the exact same focal point with the first taken under a shaft of full-sun light and the second, only minutes later, under an overcast sky. Fall witchgrass is either one species of a two-species genus (Hitchcock and Chase (1950, p. 585), a single species consisting of two varieties that comprise a monotypic genus (Gould, 1975, p. 419), or two species among several in an expanded Digitaria genus (Barkworth et al., 2003, 362-363). Classification of this species has long-been problematic. In Texas Grasses Silveus (1933, ps. 494-495) recognized L. cognatum as the only Leptoloma species in Texas and noted that it differed from Digitria species in having long-pediceled spikelets on a loose panicle in contrast to short-pediceled spikelets on thin racemes as in case of Digitaria species. In this vein, the other common name given by Silveus (1933, p. 494) for L. cognatum was diffuse crabgrass. In Botany of Western Texas (Coulter, 1891-1894, p. 508) had included fall witchgrass within an immense Panicum genus as P. autumnale closely aligned with such species as P. capilliare, P. hallii, P. virgatum, and P. bulbosum. Incidentially, the adjective "fall" almost assuredly was derived from the specific epithet autumnale. Fall witchgrass is a locally important, perennial, forage species on Texas prairies and throughout the Edwards Plateau. Beginners in these areas often confuse fall witchgrass with the annual species, called witchgrass or common witchgrass (P. capilliare). Common association of these two species is one reason for this confusion even though they are distinctly different when campared hand-in-hand. There again, Coulter (1891-1894, p. 508) aligned these two species closely so calling one for the other in understandable. Fall witchgrass (and common witchgrass for that matter) is a shortgrass or, possible interpreted as a midgrass species with panicles comprising one-third to one-half the height of the shoot. Hatch and Pluhar (1993, p. 127) described fall witchgrass as having fair feed value for both livestock and wildlife. This species is widespread in . It favors somewhat disturbed microsites and fits the classic description of an increaser. Its biological range extends from Ontario south to Florida and west to Texas northward to central Nebraska. Fall witchgrass is found in all ten vegetational (land resource) areas of Texas and in Oklahoma is absent primarily from the southeastern forest area. Crinkled or wavy margins of leaves (at least on one edge) is a distinguishing feature that is sometimes--though by no means always-- present (Hatch and Pluhar, 1993, p. 127). This feature seems to be a more widespread characterfrom southern Oklahoma through central Texas (Coffey and Stevens, 2004, p. 73) than farther to the north and east in the Tallgrass Prairie Region. Ottawa County, Oklahoma. Mid-July; full (though immature)-panicle stage.

195. Composite view- General view (under an overcast sky) of a portion of one plant of fall witchgrass showing features of leaves, culms, and panicle. This and the next two photographs showed that roughly a third to a half of total height of mature plants is in the comparatively large panicle. Ottawa County, Oklahoma. Mid-July; full (though immature)-panicle stage.

196. Sexy increaser- Detailed composite views of sexual shoots of fall witchgrass showing most of the panicle and subtending shoot (first slide) and particulars of plant features at axillary region and point of panicle emergence from boots (second slide). Fall witchgrass is an increaser on most range sites with its natural increase by both sexual and asexual reproduction with vegetative increase by both tillers and rhizomes (Hatch and Pluhar, 1993, p. 127). Although semi-cespitose in habit, fall witchgrass develops into small colonies with daughter plants (modular units) arising from the short rhizomes. The forb in the upper left of the first photograph (and also visible in the two preceding slides) was a species of tick-clover or tick-trefoil (Desmodium sp.). Ottawa County, Oklahoma. Mid-July; full (though immature)-panicle stage.

197. Big leaves on a small grass- Details of leaves, including axillary area (collar), of fall witchgrass. Ottawa County, Oklahoma. Mid-July; full (though immature)-panicle stage.

198. Maximum development- Fall witchgrass with ripening caryopses on the panicle as seen from a more-or-less topdown view. Portion of a fairly large colony with several sets of shoots that arose from short rhizomes. This was a good example of both sexual and asexual (vegetative) reproduction in this increaser species that is widely distributed across southern grasslands. Ottawa County, Oklahoma. Mid-July; full--and mature--panicle stage.

199. Paspalum plants- Two individual (two genetically unique) plants of field paspalum (Paspalum laeve) on degraded prairie in the Ozark Plateau. Individual shoots were discernible, but details of shoots were reserved for succeeding slides. These two plants provided typical examples of the habit (including the pattern of wide spacing of shoots) of this prairie and savanna grass species that usually grows to middle heights (usually not more than two and a half feet with roughly a foot and a half being more common on habitats typical of this region). This species has short rhizomes from which ascending tillers arise resulting in a somewhat at sprawling plant with widely spaced tillers. This habit was best seen in the first of these two photographs. Field paspalum is frequently found (mostly as isolated plants or a few local concentrations of plants) on oak-hickory forest-tallgrass prairie savannahs, especially in the Prairie Peninsula, in the Ozark Plateau as well as on tallgrass prairie in the Central Lowlands physiographic province. Field paspalum appears to be adapted to a wide array of microhabitats from pristine to disturbed. The two plants in these photographs were both growing on roadsides subjected to infrequent heavy mowing. They were growing about a mile distant from each other. There are a number of Paspalum species on tallgrass prairie and related mesic grasslands, but none of these are major (ie. dominant, abundant, critical or indicator) species. The paspalums are generally quite palatable, especially to cattle and horses, but they do furnish preferred forage in late maturity through dormant stages. Ottawa County, Oklahoma. Mid-July early bloom stage.

200. The hairy leaf axils of field paspalum have been described by many workers (Coulter, 1891-1894; p. 499; while others such as Hitchcock and Chase (1950, p. 615) described sheaths as "glabrous or nearly so" and included plants similar to P. laeve but having pilose sheaths as P. longipilum. Steyermark (1963, p. 202) in Missouri Flora (the definitive flora closest to location of these photographs) distinguished between P. laeve var. pilosum with "strongly hairy" sheaths from P. laeve var. laeve with glabrous or nearly glabrous" sheaths. Examples shown here were obviously P. laeve var. pilosum. In Texas, Gould (1975, ps. 526-527) recognized three varieties of P. laeve including P. laeve var.pilosum which he interpreted as synomous with P. longipilum. Barkworth et al. (2003, ps. 572-573) eliminated three varieties of P. laeve and the species, P. longipilum listed in their synonymy (Barkworth et al., 2003, p. 748) and lumped all four of these taxa under P. laeve. Another interesting feature of P. laeve is crinkled or corrugated margins (wavy edges) of leaves. This characteristic was shown in Hitchcock and Chase (1950, p. 618, Fig. 895) and Barkworth et al.(2003p. 573) This character was visible in the second and third slides in this three-slide set. In essence, the Paspalum genus has survived--as of this writing--onslaughts of cladists, but there has been considerable variation in interpretation of taxa over admittedly relatively minor features such as pubescence of leaf axils. Ottawa County, Oklahoma. Mid-July early bloom stage.

201. Lower parts- Details of basal shoots of field paspalum that was thriving on a disturbed roadside on a bottomland range site in the western Ozark Plateau. Field paspalum produces short rhizomes from which tillers arise.The first of these two slides showed several tillers with bulbous (sacculate or inflated-appearing) nodes. Such tillers frequently produce sister tillers or branches off of the parental shoot at these large, swollen-appearing nodes. Portions of two of these sister tillers (daughter shoots) were shown at close camera distance in the second slide. Ottawa County, Oklahoma. Mid-July early bloom stage.

202. Sexual reproduction from asexual reproduction- In field paspalum older tillers that are still solely in vegetative stages (ie. are asexually reproductive) often have a semi-prostrate morphology (somewhat of a creeping habit) near the ground surface that ascends at higher and younger portions (see again whole-plant shots above). This results in a shoot that resenbles a stolon off of which daughter tillers (modules of vertical shoots) develop. The secondary (daughter) tillers are considerably shorter than primary (parental) tillers and these clonal offshoots frequently produce small infloresecences that usually do not fully emerge from the boot (enveloping sheath). Two such basal inflorescences (still in the boot) on shoot branches were visible in the first photograph. The second photograph presented two secondary or daughter tillers (vertical sister shoots). The secondary shoot on the left had its nflorescence emerging from the boot (about to the final degree to which it would be expressed). The daughter tiller on the right had its inflorescence still completely encased within the boot. Pictorial representations of basal flower clusters in field paspalum were shown in line drawings in both Hitchcock and Chase (1950, p. 618, Fig. 895) and Barkworth et al. (2003, p. 573).

203. Cluster and spikelets- Details of a sexual shoot with inflorescence (first photograph) and separate branches with flowering spikelets (second and third photographs) of field paspalum on a disturbed bottomland range site in the western Ozark (Springfield) Plateau. The inflorescence of Paspalum species has been interpreted variously by different workers. Hitchcock and Chase (1950, p. 599) regarded the flower cluster as as consisting of "... one to many spikelike racemes, solitary, paired, or several to many on a common axis". Hignight et al. (1988, pos. 7, 28) provided one of the most exacting and specific categorizations of Gramineae inflorescence types ever and described the Paspalum inflorescence as a panicle of alternate spicate primary unilateral branches. Barkworth et al (2003, p. 572) interpreted the Paspalum inflorescence as a panicle of racemose (racemosely arranged) branches. In P. laeve spikelets occur singlely yet strongly overlaping in two rows along the racemose or spicate primary unilateral branch. This arrangement produces a crowded or jumbled apperance of these broad, oval flowers and grains (second slide) on the branch or rachis. The third slide showed one of the unilateral panicle branches along most of its length at peak anthesis. Anthers and stigmas were visible, at least in the photograph prior to it being scanned in an Epson Perfection scanner (slip-shod equipment which is far from perfection in regards to clear, detailed reproduction.of images). Ottawa County, Oklahoma. Mid-July early bloom stage.

204. Paspalum on the prairie- Local stands of Florida paspalum (Paspalum floridatum) on mesic tallgrass prairie in the western Ozark Plateau. Florida paspalum is typically the largest of the Paspalum species on tallgrass prairie and savannas of tallgrass prairie and oak-hickory forests. On coastal tallgrass prairie such as those of the Gulf coast there are Paspalum speaacies that grow to larger size, but on prairies of the continental interior Florida paspalum generally takes the size prize. It is also the single most important Paspalum species in the Tallgrass Prairie Region. For example, it was the only Paspalum species included in Field Guide to Oklahoma Plants by Tyrl et al. (2008). Leiithead et al. (1971, p. 137) and Tyrl et al. (2008, p. 147) reported that Florida paspalum is quite palatable, especially to cattle, though forage quality declines rapidly in this succulent species after dormancy (a situation that applies to all Paspalum species). Given this palatability, Florida paspalum is apt to be heavily utilized so as to benefit from periodic deferment during the growing season every second or third year (Leithead et al., 1971, p. 137). The Paspalum species are similar in general apperarnce and, in case of some of the native species, in morphology of inflorescences and details of spikelets. The color photographs in Coffey and Stevens (2004, ps. 89- 94) are useful in helping beginners (sometimes even "experts") distingush among Paspalum species common in southern Oklahoma and northcentral Texas. Barkworth et al. (2003, ps. ) is the new definitive source for North American paspalums. Ottawa County, Oklahoma. Late July; peak standing crop and anthesis.

205. Study in grass sex- Anthesis in Florida paspalum with anthers and stigmas exerted simultaneously. Florida paspalum is one of the best examples of a native prairie grass in which to show the sex organs because typicall both female and male organs are expressed at the same time. Ottawa County, Oklahoma. Late July; peak anthesis.

206. Done for the year- Dead shoots of knotroot bristlegrass (Setaria geniculata) shortly after this plant entered dormancy. Knotroot bristlegrass is a widely distributed panicoid grass that prefers mosit habitats such as those along streams or seeps (Gould, 1975, p. 546). It is a common, characteristic species on the Fort Worth or Grand Prairie in northcentral Texas especially on disturbed sites (Tyrl et al., 2008, p. 161) such as overgrazed ranges where the scarcity of climax little bluestem and Indiangrass results in reduced competition. Erath County, Texas. November, grain-shatter at dormant stage.

207. Opposite ends- Two dense panicles (first photograph) and the characteristic swollen nodes on short rhizomes (second sphotgraph) of knotroot bristlegrass. This species has one of the more descriptive common names of grasses, but one does have to pull a specimen up to see the source. Erath County, Texas. November, grain-shatter at dormant stage.

208. New cords and old cords- Prairie cordgrass growing in the Platte River Valley with green shoots of the current growing season along with dead shoots . from last summer's growing season. Adams County, Colorado. Early July; pre-boot stage of phenology.

209. Collar and ligule- Two views of the axillary (leaf axil) portion of shoots of prairie cordgrass. Serious and comprehensive presentations of grass features should include (when possible) details of collar, ligule, sheath, blade, node, internode, veination, and so on. Adams County, Colorado. Early July; pre-boot stage of phenology.

210. Bottom portions- Lower portions of shoots of prairie coregrass. The first slide was of shoot bases of both dead (from the previous warm-growing season) and live (current spring andsummer-growing season) Spartina pectinata shoots growing on the floodplain of Platte River. The second photograph included rootcrowns and a short (young) rhizome of this strongly rhizomatous, native, eragrostoid grass. This C4 grass reproduces asexually via rhizomes and tillers so that it is often found large and (apparently or presumedly) clonal stands. Prairie cordgrass usually forms relatively extensive colonies in more mesic areas such as prairie sloughs (see above where vegetation of range types was featured). Preference of prairie cordgrass for sloughs is basis for another common name of sloughgrass (Shaw, 20008, p. 183), but this common name should be reserved for Beckmannia syzigachne which has the preferred (published) common name of American sloughgrass. This latter species frequently occurs sympatrically with prairie cordgrass, but is an annual and does not form expansive colonies. Adams County, Colorado. Early July; pre-boot stage of phenology.

211. Crowned up and starting to crawl- Rootcrown and rhizome of prairie cordgrass on the floodplain of Platte River. These two photographs presented the the scaly phytomers (tough bract [sheath]-covered node-internode units) of shoots--both tillers and rhizomes--at or just below soil surface (O horizon). The first slide showed a rhizome growing outward from two tillers (top of slide) with a new tiller (daughter or sister shoot) arising from the rhizome and then either the rhizome growing still outward or, alternatively, the daughter shoot producing a rhizome (depending on interpretation). The second slide showed the large number of basal phytomers of a tiller and another tiller (sister or clonal unit; another module) from which a short (immature) rhizome was growing. The rhizome is also a sister or sister shoot although horizontal and completely subterranean in contrast to daughter tillers. The vertical tillers are intravaginated shoots (ie. the outward growing shoot does not perforate or pierce the invaginating sheath) whereas the horizontal rhizomes are extravagated shoots which do perforate of burst through the enveloping sheath(s). Observe the organic matter-rich alluvium of the Platte River floodplain that served as the natural medium for growth of this rank, perennial, tallgrass species. Adams County, Colorado. Early July; pre-boot stage of phenology.

212. Crawling quite a way- A long rhizome of prairie cordgrass against a backdrop of tillers growing in the fertile alluvium of the Platte River floodplain. Obviously, this specimen (like those in preceding photographs) was dug out and exposed for photographic purposes. Adams County, Colorado. Early July; pre-boot stage of phenology.

213. Cluster of racemes- Inflorescences of prairie cordgrass growing on a plains cottonwood-mixed prairie savanna that developed along an intermittent stream in the Colorado Piedmont. The inflorescences of Spartina species are confusing. Traditionally the branches with units of spikelets aligned along them were interpreted as racemes or, maybe, spicate racemes, but as these spikelets are sessile this view was at least somewhat inconsistent with standard definitions. Hitchcock and Chase (1950, p. 508) regarded the Spartina flower cluster as being made up of "... appressed or sometimes spreading spikes racemose on the main axis". Later, more detailed (sophisticated) descriptions of grass inflorescences recognized specialized panicle inflorescences (Highnight et al., 1988). Using this scheme, Stubbendieck et al. (1992, p. 107) described the inflorescence of prairie cordgrass as a "panicle of 6-40 spicate pirmary unilateral branches". Shaw (2008, p. 183) described the inflorescence of prairie cordgrass as a panicle with racemosely arranged, appressed, alternate branches. East Bijou Creek, Elbert County, Colorado. Mid-July; peak-bloom stage (full anthesis).

214. Anthers, anybody?- Two close-up views of racemes in full anthesis on prairie cordgrass. The first photograph presented a standard side-view perspective of the raceme while still on the plant whereas the second photograph gave a ventral view of a raceme that was possible only by removal from the shoot. (The latter was laid on cattle dung in situ on a floodplain cottonwood-mixed prairie range that developed along an intermittent stream.) There are often so many spikelets (one-flowered and sessile) on the racemose, unilateral branches that these floral units overlap (ie. imbricate spikelets). The Spartina species often produce some of the greatest number of anthers of any grass genera native to North America which in combination with imbricate spikelets produces an extremely showy array of exerted anthers. That characteristic was seen in these photographs. East Bijou Creek, Elbert County, Colorado. Mid-July; peak-bloom stage (full anthesis).

Sporobolus complex- A group of closely Sporobolus species, varieties of one species, or varieties of two or more species with a wide biological range have long been recognized as important on tallgrass prairie. Tall dropseed, as a catchall common name, has been treated as including up to three varieties of the species S. asper or, as a synonym, S. compositus. . The specific epithet asper means "tall". More recently it was discovered that the author of S. asper used as basis of his scientific name an earlier designation that by international botanical rules was "illegitimate" (Tyrl et al., 2008, p. 173) so that the binomial now regarded as correct is S. compositus. If this confusion was not enough there has been the long-standing view that S. compositus (S. asper) is so "morphologically variable" that perhaps it is "conspecific" with S. clandestinus (Tyrl et al., 2008, p. 173). All told this mess amounts to what taxonomists regard as a "complex". The Sporobolus compositus complex is a difficult assemblage of forms, perhaps affected by their primarily autogamous breeding" which is largely responsible for these plants being able to "perpetuate unique character conbinations" (Barkworth et al., 2003, ps. 121-122 ). Autogamy refers to self-fertilization within a flower. Much of this autogamy occurs in cleistogamous spikelets, those having "self-contained" fertilization (Gould, 1975, ps. 303, 631). By whatever name(s)--at species or variety level--there are at least three taxonomic forms under the scientific "handle" for a generic tall dropseed. Presented immediately below were two of these: 1) "regular" tall dropseed (S. compositus var. compositus= S. asper var. asper) and 2) meadow dropseed (S. compositus var. drummondii = S. asper var. drummondii).

215. Tall tops- Sexual shoots of tall tropseed (Sporobolus asper var. asper= S. compositus var. compositus) with spikelike panicles retained within their boots (enclosing sheath). The "dead giveaway" of tall dropseed is the "buggy whip"-appearing extension of the panicle-enclosing sheath. Unfortunately this very distinctive feature was often missed by botanical artists so that illustrations of this species (variety) frequently did not show the most foolproof feature for field identification. For example, this 'buggy whip" flagellate extension of the enclosing sheath was not shown in Hitchcock and Chase (1952, p. 416) nor in the popular Phillips range and pasture plant book (Phillips Petroleum Company, 1963; Fort Hays State University, 2006, p. 65). One can pretend that the prolonged extension of the sheath had been grazed or weathered off in these publications. Fortunately, more accurate renditions were shown in Barkworth et al. ( , p. ) and Tyrl et al. (2008, p. 172). Ottawa County, Oklahoma.

216. Grraminoid buggy whips- Two inflorescences of tall dropseed growing on degraded tallgrass prairie ranges: the first from the Ozark Plateau and the second from the West Cross Timbers. The flagellate extension of the sheath that encloses the contracted panicle of Sporobolus asper var. asper (= S. compositus var. compositus) is a very distinctive feature of this variety. This morphological characteristic tends to be more commonplace in wet years. Both of these sexual shoots were produced in a widespread, ususually wet late summer through autumn First slide: Ottawa County, Oklahoma. Mid-December. Second slide: Erath County, Texas, Early November.

217. Pushed and bleached out- Basal part of a sun-faded spicate (contracted) panicle of tall dropseed within its enveloping sheath. This partial exertion (or, same difference, retention) of the panicle within the boot is a feature of several of the Sporobolus species. The fruit (achenes in strict definition) had shed from this contracted panicle which was the same one presented in the immediately preceding photograph. Erath County, Texas, Early November, fruit-shatter stage.

218. Bunches on the meadow- Several plants of meadow dropseed (Sporobolus asper var. drummondii= A. compositus var. drummondii) Although perennial dropseeds are generally increaser species, meadow or Drummond's dropseed is a member of the climax grassland community in some types of tallgrass prairie. Such is the case for specific range sites of the Grand or Fort Worth Prairie in northcentral Texas. There are several varieties (or whatever taxa is used by this or that authority) that comprise a Sprorbolus asper (S. compositus) complex. At this stage of development all leaf and most of culm growth had already taken place. Flowering would not be until late summer. Erath County, Texas. Early May.

219. Short with a puffy boot- Puffsheth dropseed (Sporobolus neglectus) on a shallow habitat of a deteriorated range. Typical example of the small, annual species. Most of the Sporobolus species are perennials that are most abundant on abused ranges, especially overagrazed pastures. The same is true of puffsheath dropseed, but it is even more common on abandoned fields in early seral stages (ie. go-back land or old fields). Puffsheath dropseed is not so much a pioneer or initial colonizing species as it a second-stage invader although it is commonly associated with colonizing annual grasses and forbs. Tyrl et al. (2008, p. 177) explained that S. neglectus along with two or more other Sporobolus species constitute an "interbreeding comples". This integrading group has been intrerpreted variously by different authorities, but to this authors eyes S. neglectus is noticably distinct due to its diminutive size and short inflorescences. Tarleton State University Hunewell Ranch, Erath County, Texas. Late October, fruit-ripe phenological stage.

220. Puffed but not out- Details of sexual shoots of puffsheath dropseed. The small, contracted panicles of this annual species were still largely retained within the boot, the enveloping sheath surrounding the flower cluster. Thus, basis of the adjective of the common name. It was remarked above that this partial retention of the panicle is a characteristic of many of the Sporobolus species. Tarleton State University Hunewell Ranch, Erath County, Texas. Late October, fruit-ripe phenological stage.

221. Heavy infestation of common grassbur or sandburr- Cenchrus incertus is one of the most abundant grasses in Texas and south into Mexico. It is a weed of overgrazed pastures and ranges, suburban yards, and abandoned city lots (in short, on any disturbed soil surface). It is usually a short-lived perennial or, sometimes, an annual. If ever there was a meek to inherit the earth of Texas it would be this spiny weed, though for that reason some would make the case it most certainly is not meek. (In fact, it is a pretty good match for the personality of many Texans who fashion themselves as half-hoss and half-gater just not quite wild enough to eat grassburrs.) Erath County, Texas. July.

222. Involucres on grass or sand burr- The sharp-spined spikelets on this "black sheep" of the prairies are a real pain in the you-know-what (and about e9erywhere else). Grass burrs are a textbook example of zoochory, dispersal of plant disseminules by animals, including rangemen.

223. Another native, cool-season, prairie grass- Winter bentgrass (Agrostis hyemalis) on moist to wet local habitat on tallgrass prairie in western Ozark Plateau. This is a minor species other than in local mesic microsites. It was included here to give an idea as to the great species diversity of tallgrass prairie. Even in southern areas native, winter, grasses grow alongside warm-season, perennial, panicoid grasses. Winter bentgrass has generally been regarded as a short-lived perennial (Gould, 1975, p. 140). It grows on both interior and coastal tallgrass prairie. Ottawa County, Oklahoma. Late May, maturing grain to early grain-ripe phenological stages.

224. Dainty panicles on tallgrass prairie- Panicles and spikelets of winter bentgrass. These flowers were on some of the same plants shown in the immediately preceding slide. Ottawa County, Oklahoma. Late May, early grain-ripe stage..

225. White tridens (Tridens albescens)- White tridens is another cool-season native grass common on prairies and prairie-forest edges, especially in local moist habitats such as depressions and edges of imponded water. Unlike the preceding wildryes and Junegrass that usually respond as decreasers, white tridens is typically an increaser in response to disturbances like overgrazing. This species is particularily common and conspicuous on mesic Texas grasslands including the Grand, Blackland, Fayette, and North Central Prairies as well as adjoining savannahs like the Cross Timbers and Post Oak Savanna. White tridens often forms large stands on depressions, along banks of ephemeral stream and ponds, and other places where soil moisture can accumulate or even inundate the land surface for brief periods. By nature of its preference for a wet rooting medium white triden often grows on clay or generally heavy textured soils. The stand of white tridens seen here was growing on a depression just upslope from a shallow stream in the Western Cross Timbers area, a made-to-order home for white tridens. Lee Ray Ranch, Stephens County, Texas. May (and a wet one).

226. Customary stance- Typical habit of white tridens. This cespitose grass of intermediate size (a mid-grass species) usually has a sprawling or "spread-open" physical appearance as shoots come off from the basal part of the plant at angles less than vertical. Lee Ray Ranch, Stephens County, Texas. May

227. Panicles of white tridens- The "tight-packed" spikelets of numerous florets on short pedicels along a central floral axis make up a contracted panicle inflorescence type in white tridens. The panicles in this frame are just past anthesis with the tiny grains in the milk to soft dough stage. Lee Ray Ranch, Stephens County, Texas. May.

228. Tinge of purple in white tridens- Viewed at close distance the coloration pattern of white tridens is quite pleasing with traces of dark blue, purple, or bluish red splotched across the lemmas.Some of the other Tridens species have similar colors. Hitchcock and Chase (1950, p.216) described white triden spikelets as having eight to twelve florets whereas Gould (1975, p. 209) put the number of flowers at four to eleven. Lee Ray Ranch, Stephens County, Texas. May.

229. Tumblegrass (Schedonnardus paniculatus)- This distinctive grass is a native perennial that is regarded as an invader or, depending on range site, an increaser at best. It is a characteristic member of early seral stage communities and thus is more common on deteriorated ranges, sacrifice areas, etc. It is one of the earlier perennials to become established on go-back land (ie. a component of seral stages that are slightly more advanced than the pioneer stage). Consistent with early seral species, tumblegrass has poor forage value. Texas A&M University Agricultural Experiment Station (Stephenville branch station), Erath County, Texas.April.

230. Inflorescences of tumblegrass- The inflorescence of tumblegrass is a raceme which is characteristic of the Chlorideae tribe of which this weedy grass is a member. This "seedhead" is quite distinctive. It is of a relatively large size and frequently breaks off from the basal part of the plant and rolls along the ground surface much like a tumbleweed (hence it's common name). Erath County, Texas. June.

231. Local stand of tumble windmillgrass (Chloris verticillata)- This dense population of tumble windmillgrass with a typical pattern of inflorescences was growing on a disturbed site-- the most common category of habitat for the Chloris species-- in the Cross Timbers of northcentral Texas. Chloris species are not major forage producers nor are they usually climax species. Some Chloris species are interpreted as ecological invaders while others are classified as increasers. Erath County, Texas. July.

232. Inflorescences of tumble windmillgrass- The inflorescences of the Chloris species has been a subject of semantical debate among agrostologists. Traditionally the inflorescence type of Chloris (and some related genera) was called a raceme (an unbranched inflorescence with spikelets pedicellate-- pediceled-- on the rachis) as typical for members of the Chlorideae tribe (eg. Bouteloua, Buchloe, Spartina). In this view the individual racemes are each separate inflorescences. Alternatively the individual racemes can be viewed as part of a general compound raceme or, as another alternative, as a number of one-sided spikes that are either solidary, digitate, or racemose on a main axis. More recently, some workers came to interprete the entire inflorescence as a panicle of of verticillate (arranged in whorls) spicate (ie. spikelike) primary unilateral (one-sided) branches. A nice debate about a genus of relatively little consequence relative to so many others. In Cross Timbers vegetation, Erath County, Texas. July.

233. Perennial threeawn (Aristida purpurea) on deteriorated tallgrass prairie range - What is most likely "purple threeawn" was thriving and adding spring color to an area heavily impacted by motor vehicle traffic.This was typical "prime habitat" for threeawns which are some of the most abundant perennial grasses on the most abused areas (eg. sacrifice areas, "stomp lots", ranch roads). The other most common species on this deteriorated site was the naturalized alien, rescue bromegrass. The threeawns or "wiregrasses", as they referred to in the southeastern pine forests, are typically classified successionally as invaders. As a general rule Aristida species are characteristic of vegetation at some stage of retrogression and therefore indicator plants of deteriorated ranges (or regenerating pine forests in less advanced seral stages). One of the general features of species responding as ecological invaders is low feed value. This is most pronounced in annual species, but some perennial species (in fact, entire genera) have this feature. Aristida is one of these (with some rare exceptions). In fact, there is a high proportion of Aristida species that are annuals, even short-lived annuals known as ephemerals. As if all this were not enough, Aristida includes several taxa that are a taxonomist's worst nightmare. Some of these integrade or hybridize (or whatever grass plants do "to make life harder for agrostologists") with the result that various authorities have interpreted such taxa as species, subspecies, and varieties in different ways which has led to confusion and frustration (especially because the taxonomic arrangements seem to change with every new treatment of Aristida). The term that has often been invoked in describing and arranging these taxa into groups is "complex". An example of such includes such closely related and intergrading threeawns as red threeawn (A. longiseta), purple threeawn (A. purpurea), wright threeawn (A. wrightii), roemer threeawn (A. roemeriana), and a few others depending on which of these species grow in association in any given geographic location. In Texas the complex is often known simply as "perennial threeawn" or, sometimes, "purple threeawn" after A. purpurea which is viewed as the species, with what have previously been interpreted as separate species ranked as varieties therein. Thus "perennial or purple threeawn" includes A. purpurea var. longiseta, A. purpurea var. purpurea, and A. purpurea var. wrightii. Other agrostologists still designate these threeawns known by the various common names as separate species. There was bound to be some sort of lesson in this example of "where the experts cannot agree".

234. Examples of perennial threeawn- These photographs displayed some individual plants that were typical of "perennial threeawn" (A. purpurea complex) in the Cross Timbers and Prairies vegetational area of northcentral Texas. Aristida species are strictly cespitose (= bunchgrass or tufted) species that produce only intravginated shoots (tillers) and no extravaginated shoots (stolons and rhizomes). There are some species in Sporobolus, Muhlenbergia, Festuca, and Andropogon (among others) that also have the cespitose habit, but in North America Aristida is probably the most extreme case of this. The tufted habit was very obvious in these examples.

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235. Old-field threeawn (Aristida oligantha) on an old-field- Dog-hair stand of the annual old-field threeawn or annual wiregrass on small parcel of go-back land of Fort Worth or Grand Prairie in noethcentral Texas. Most stands of this r-selected or pioneer species are not this dense, but this demonstrated the physiogonomy and structure of such populations of this species that was performing the essential function of protecting soil made bare by the Clementsian process of denudation (in this case, plowing). Tarleton State University Hunewell Ranch, Erath County, Texas. October.

236. Plants of old-field threeawn- Numerous individual plants (genotypically unique individuals) of the annual Aristida oligantha. Zoom-in view on some of the plants presented in the immediately preceding slide to show habit of this pioneer species. While old-field threeawn is an ecological invader it is valuabel-- often locally essential to prevent soil erosion and facilitate modifications in its habitat that prepare the way for species of higher successional order. This improvement in the environment (often edaphic modifictions like addition of organic matter with this species) is the Clementsian process of reaction or, in contemporaty terminology (and same thing), facilitation. Tarleton State University Hunewell Ranch, Erath County, Texas. October.

237. Old-field threeawn (A. oligantha)- There are many annual Aristida species. Some of these are, as noted above, even epemeral (ie. the so-called "six-week grasses"). Other annual species take somewhat longer to complete their annual life cycle, but are nonetheless very opportunistic. Old-field threeawn was appropriately named for its common occurrence as a pioneer species on abandoned farm fields ("old-fields"). In fact, it is probably the most common native annual grass colonizing disturbed areas like old-fields in the the central prairies prairies region (Central Lowlands and eastern Great Plains). Old-field threeawn is an example of an r-selected species (those in which natural selection has been for traits which maximize the intrinsic rate of increase such as to allow members to rapidly colonize newly or recently created habitats) covered in standard Ecology texts. For practical purposes the annual threeawns have no forage value for livestock or big game species. The specimen, the sporophytic generation, shown here had just completed its short life cycle tht streatched from early spring or late winter to early summer. Erath County, Texas. July.

238. Little 'un on the prairies, plains, and Cross Timbers (and a lot of other range to boot)- Two plants of little barley (Hordeum pusillum), a native, cool-season annual grass locally common on tallgrass prairie and tallgrass savannah. This species is found frequently on the prairies of northcentral Texas and adjoining Cross Timbers (where these plants were photographed) and extending westward throughout the Great Plains clear across to British Columbia and northeastward to Maine and down south to Florida. Given its tremendous species range, nativeness, annual life cycle, and local commonness little barley was included on the list of 200 plant species for the Society for Range Management (SRM) International Range Plant Identification Contest (Stubbendieck et al., 1992). Little barley is also included in the Texas Intercollegiate Range Plant Identifiction Contest sponsored by the Texas Section, SRM as well as on both the range plant list for Texas 4H and Future Farmers of America Contests (Hatch and Pluhar, 1993). Little barley is classified as an invader what with being an annual--even if a native one--on range where almost every one of the dominant plant species is a perennial. Little barley typically invades disturbed areas such as overgrazed ranges and "go-back land" where it undoubtedly provides some soil protection against the agents of soil erosion. A concise yet informative description of little barley, including practical aspects, was in both editions of Pasture & Range Plants (Phillips Petroleum Company, 1963; Nicholson, 2006), although the author of Range Types disagreed that little barley is "essentially worthless" for livestock and wildlife. In point of fact, this short-statured species sometimes provides forage for cattle, sheep, horses, and deer. On dormant tallgrass prairie where the herbage of bluestems, Indiangrass, sideoats grama, etc. is weathered after three months of dormancy this C3, festucoid grass frequently provides some green feed--the quantity varying substantially--thart is richer in carotene, higher in crude protein, and better in flavor than that in the herbage of the decreasers and increasers. This could be especially to the smaller ruminant species. Erath County, Texas. March; just past anthesis to early milk stage.

239. Basal properties- Shoot bases of little barley at phenology ranging from anthesis to milk stage. Erath County, Texas. March.

240. Barley spiked- Spike of little barley (H. pusillum) growing in West Cross Timbers. Spikelets of little barley are one-flowered (one floret per spikelet). Erath County, Texas. March; milk stage of phenololgy.

Grasslike Plants of Tallgrass Prairie

Tallgrass prairie includes or, at least, is adjacent to floristically associated wetlands, most of which are marshes. Most rangemen would probably interpret such marshes as part (a range site) of the overall grassland community, most logically as another range cover type within the general tallgrass prairie. In some locations these wetlands are formed from springs that drain directly into nearby creeks which are corridors of the Eastern Deciduous Forest, especially forms of the Oak-Hickory Association. In some such range vegetation, marshes abutt both tallgrass prairie bottomland and bottomland forest dominated by species like sycamore (Plantanus occidentalis), (Acer segundo), and ashes (Fraxinus spp.). In these marsh habitats there are various species of bulrush or tules (Scirpus spp.). A few tules or bulrushes were included below. Taxonomy of these remians a "mess". For example, the most useful manual for the tallgrass prairie marshes of eastern Oklahoma and Kansas was the Missouri flora. Treatment of Scirpus spp. changed dramatically from the original Flora of Missouri (Steyermark, 1963) and Steyermark's Flora of Missouri (Yatskievych, 1999). This included changing some former Scirpus species to Schoenoplectus species which was consistent with some other treatments (eg. Diggs et al. (1999). This author did know if the dead Dr. Steyermark would have approved of these change, but the non-taxonomist author of this publication did not. Scientific names reflect that opinion. Other range plants common in-- and, indeed, often comprising dominants and associates of-- marshes are various grasslike-plants especially species of. Carex, Cyperus (Cyperaceae), Juncus (Juncaceae), and Typha (Typhaceae). Several of these lesser-known, but often locally important grasslike-plant species were included below.

241. Soft rush (Juncus effusus)- This is a widely distributed rush in the tallgrass prairie region. It was found to be an important forage species to the American muskrat (Ondatra zibethicus).  Craig County, Oklahoma. June

242. Grasslike neighbors-Two species of grasslike plants on a wet tallgrass prairie in the western Ozark Plateau. Short-fruit or whiteroot rush (J. brachycarpus) and Frank's caric sedge (Carex franki) growing in association on a local ponded area of big bluestem-dominated prairie. The rush is in front of the sedge in both of these slides. Ottawa County, Oklahoma. July; early fruit-ripening stage.

243. Short-fruit or whiteroot rush (J. brachycarpus)- On a local depression with periodically ponded water and slowly drained soil this rush was growing in association with beaked panicum, broomsedge bluestem, and several species of caric sedges including fox sedge (Carex vulpinoidea) which was presented below. Whiteroot rush is not usually a common species, but it is frrequently present as individual to small numbers of plants in particular on areas of wet prairie. Palatability of whiteroot rush was unknown to this author, but it is a common observtion that generally speaking none of the Juncus species are very palatable. Economic value of Juncus species is undoubtedly in soil protection and improvement through addition of organic matter. Ottawa County, Oklahoma. July; early fruit-ripening stage.

244. A distinctive topknot- The spherical inflorescence of whiteroot or short-fruit rush makes identification of this Juncus species a fairly straightforward endeavor. Ottawa County, Oklahoma. July; early fruit-ripening stage.

245. Inland or interior rush (Juncus interior)- These two specimens were growing on the same wet paririe (though a different local depression) as many of the other grasslike plants presented in this section. This degraded tallgrass wet prairie was in the western Ozark Plateau. The first specimen (first photograph) was in mid-flowering (just post-anthesis with some stigmas still alive). Second specimen (second photograph) was at fruit-ripe to fruit shatter stage. An interesting and, to a plant photographer, obvious variable about this species (at least at this location) was the variation in time of flowering and fruitset of different plants growing within a few feet of each other. Phenological stages were similar overall, however. The less mature plant (first slide) was photographed two weeks earlier than the more mature plant (second slide). Both plants fulfilled their role: they recombined genes, yielded fruit, and helped perpetuate their race. Ottawa County, Oklahoma. Mid-June and late June.

246. Inland inflorescences and fruit- Panicle and fruit of inland or interior rush. The fruit of Juncus is a locusidal capsule). These organs were on plants growing on a small derpression--nothing much but a mud hole--on wet prairie. Ottawa County, Oklahoma. Late May, fruit-ripening (just pre-maturity) stage.

247. Fox caric sedge (Carex vulpinoidea)- Fox sedge is but one of numerous carices found in wet prairies and marshes of the central tallgrass prairie and prairie peninsula region in the interior of North America. Ottawa County, Oklahoma. June.

248. The distinctive inflorescence of fox sedge- This is a common wetland species in the marshes and poorly drained grasslands of the Cherokee Prairie and Ozark Plateau. This one was on a wet prairie used as a hay meadow, but at the edge where it was too wet for haying equipment. Ottawa County, Oklahoma. June.

249. Details of fox caric sedge inflorescence- These two inflorescences of Carex vulpinoidea showed the namesake and unmistakenable taillike (and extremely long) bract at base of the flower cluster. Fruit (achenes) on these two sexual shoots were approaching maturity and the shatter stage. These shoots were on the same wet prairie hay meadow as those presented immediately above. Ottawa County, Oklahoma. Mid-June.

250. Troublesome sedge (Carex molesta)- The carices are a troublesome genus with many "look-alikes" and positive identifiction problematic, often unless plants can be collected at various stages of phenological development. Troublesome sedge is one of numerous carices that is adapted to various habitats varying from nearly aquatic to dry prairies and open deciduous forests. This "nifty" local population was in the western Ozark mountains of the Prairie Peninsula on a tallgrass prairie located between oak-hickory forest vegeetation. Climax dominants were big bluestem and Indiangrass, but the varied immediate range plant plant community also included the invaders broomsedge and Baldwin ironweed (Vernonic baldwini). Ottawa County, Oklahoma. June, hard-seed phenological stage

251. Inflorescences of troublesome sedge- This Carex species is in the subgenus Vignea the species of which are characterized by having two stigmas and perfect (bisexual) flowers in contrast to subgenus Eucarex whose species have three stigmas and separate staminate and pistillate flowers (unisexual). Though not of high palatability or high on animals' "preference list" carices like troublesome sedge are consumed by livestock and wildlife like deer. All the grasslike plants are valuable for soil protection and formation. Ottawa County, Oklahoma. June, hard-seed phenololgical stage.

252. Another caric sdege of wet prairie- Bicknell's caric sedge (Carex bicknellii) growing on the marsh-like drainage of a wet prairie used as a hay meadow. This was the same portion of meadow on which fox sedge (above) and Frank's sedge (below) were also important local species. Ottawa County, Oklahoma. Late May, early fruit development stage.

253. Distinctive top- Flower cluster and individual floral units of Bicknell's sedge. This is another Carex in the Vignea subgenus. Ottawa County, Oklahoma. Mid-June, ripening to mature fruit stage.

254. A caric sedge of wet prairie- Frank's caric sedge (Carex frankii) in extreme western Ozark Plateau. Several other Carex species, including C. vulpinoidea, fox sedge (shown above), and C. bicknellii , Bicknell's sedge (shown below), were growing on this wet prairie that was used as a hay meadow. Ottawa County, Oklahoma. Mid-July.

255. Boy groups and girl groups- on wet prairie- Close-in views of inflorescences of Frank's caric sedge showing both pistillate (lower) and staminate (upper) flower clusters. This Carex species is in the Eucarex subgenus which was described in the caption of troublesome sedge above.

256. In winter and in summer- Another caric sedge of moist, though not wet, prairie of tallgrass prairie in the Ozark Plateau and Cherokee Prairie was Levenworth sedge (Caraex levenworthii). It was shown here in its apparel of winter (first photograph) and of spring-summer (second photograph). Note that in its winter turf Levenworth sedge was beginning to green up by early winter. Obviously this is a cool-season species (at least in this area). Ottawa County, Oklahoma. First slide, early December (just breaking dormancy); second slide, mid-June.

257. Another distinctive cluster- Inflorescences of Levenworth's caric sedge. These two flower clusters were on plants shown in the summer sward of this species (second slide immediately above). This species of Carex is in subgenus Vignea. Ottawa County, Oklahoma. Mid-June, early fruit development..

258. Green flatsedge (sometimes written as flat sedge) or umbrella sedge (Cyperus virens= C. pseudovegetus)- This is typically an aquatic plant or hydrophyte that is one of the more common Cyperus species on wet to moist soil of the tallgrass prairie, openings in the western oak-hickory forest, and the prairie peninsula ecotone between these two major range communities. That it is readily eaten by cattle was indicated by the fact that in a pasture adjoining the prairie hay meadow in which this photograph was taken no inflorescences on shoots of umbrella sedge could be found. Ottawa County, Oklahoma. June.

259. Manyflowered unbrella sedge (Cyperus lancastriensis)- This species or flat or umbrella sedge grows on various local habitats though it usually pre8ers moister soils. This specimen was growing on an upland slope at edge of prairie and oak-hickory forest. Many of the Cyperus species resemble each other closly in morphology so that identification to species level is difficult. Many species display considerable variability in features in infloresence, spikelet and floret. Frequently there is intergrading and hyrbridization among similar species. Treatment frequently varies greatly among authorities. Ottawa County, Oklahoma. July; pre-bloom phenological stage.

260. Flowering in manyflowered umbrella sedge- Inflorescences on a plant of manyflowered unbrella sedge growing in close proximity to the individual shown in the preceding slide. This plant was more advanced phenologically and had fully exerted anthers and stigmas. Cyperus species are not as difficult to identify accurately as are many Carex species, but both are difficult enough. Much of this difficulty is due to hybrids of closely related species. By way of example, the plant shown in this and the plant presented in the precding slide fit the description and were in the near vacinity of the hybrid specimen described by Steyermark (1963, p. 267) as that between C. lancastriensis and C. filiculmis, C. ovularis, or C. strigosus. Lower spikelets of the plant presented here did not have its lowest spikelets reflexed in the manner typical for C. lancastriensis, but it clearly was of this species. An example of differing treatments of Cyperus species among authors was provided by this case. In the expanded Steyermark's Flora of Missouri (Yatskievych, 1999, p. 372) did not allude to the hybrid that Steyermark himself (Steyermark, 1963, p. 267) gave a detailed description of. Ottawa County, Oklahoma. July; anthesis.

261. Sand spikerush (Eleocharis montevidensis)- Eleocharis is another genus in the Cyperaceae. There are several spikerush species on tallgrass prairie. Sand spikerush is a represnetative species from the Grand Prairie of northcentral Texas. It is typically a short-statured plant. For example, the shoots in these photographs varied from six to nine inches in height. Tarleton State University Hunewell Ranch, Erath County, Texas. March (peak anthesis).

262. Woolgrass (Scirpus cyperinus)- The example of this distinctive Scirpus species was growing in a water-filled deep depresion on a virgin hay meadow in the Blackland Prairie vegetational area of northcentral Texas. Grayson County, Texas. June, soft-grain stage.

263. Pendant bulrush (Scirpus pendulus)- Typical plant of pedant bulrush growing in a roadside ditch on wet prairie in the eastern portion of Cherokee Prairie, a major section of the Central Lowlands physiographic province. Craig County, Oklahoma. Late May.

264. Pendulus clusters- Sexual shoot with numerous panicles (first slide) and two units or clusters of spikelets off of one panicle (second slide) of pendant bulrush on wet prairie. The shoot with its entire panicle (and with numerous units or clusters) was one of the tillers on the plant presented in the immediately preceding photograph (Craig County, Oklahoma). The panicle units had spikelets, the distal most floral units of which were in anthesis (Ottawa County, Oklahoma). Both photographs taken in Late May.

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265. Great bulrush, softstem bulrush, or giant tule (S. validus= S. tabernaemontani). The example of this species was growing in a slough at edge of a tallgrass bottomland prairie just above Lost Creek in western portion of the Ozark Highlands (Mountains). According to Steyermark (1963) this species is the common bulrush of the Ozark Plateau. The terminal end of the culm that extended beyond the inflorescence in this specimen closely resenbled that of S. acutus, but this latter species does not occur in within 150 miles or more of this location (Steyermark, 1963, p. 292-293). Ottawa County, Oklahoma. July, hard dough-grain stage.

266. Common bulrush (S. atrovirens)- This Scirpus species is probably the most common one in prairies of Missouri and eastern Kansas and Oklahoma. S. atrovirens is an extremely variable species. Steyermark (1963, p. 296) divided it into three varieties which Yatskievych (1999, ps. 429-432) elevated to three species. These plants were growing in a tallgrass prairie slough that drained into Lost Creek at the western edge of the Ozark Plateau. Ottawa County, Oklahoma. June, hard dough-grain stage

267. Common or broad-leaf cattail (Typha latifolia)- This is one of three Typha species (as commonly interpreted by most authors) found on wetlands of the central grasslands of North America. Along with members of the rush and sedge family (and some minor ones like the burr-reed family) cattails constitute the groups of range plants commonly known as grass-like plants. This group or category of range plants carries no official taxonomic recognition in the lexicon of Plant Systematics, but is used primarily by rangemen and foresters to designate those species of monocots which do not produce flowers with showy or conspicuous petals and yet which are not grasses. These include several orders of monocotyledons. Vascular spore-bearing plants such as the horsetails or scouring rushes (Equisetum spp.) and monocots with obvious petals (eg. the orchids) are classified as forbs by natural resource practitioners like rangemen and wildlifers. foresters. The grasses and grass-like plants are, in turn, known collectively as graminoids. Cattails are some of the most common and conspicuous species of grass-like plants on both marshes and wet microsites on grasslands across North America. Cattails grow not only in natural depressions like potholes and wet draws but also around man-made structures that impond water suc