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).

7. 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).

8. 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).

9. 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).

10. 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).

11. 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).

12. 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).

13. 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).

14. 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).

15. 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.

16. 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).

17. 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).

18. 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.

19. 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).

20. 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).

21. 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).

22. 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).

23. 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).

24. 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).

Great Plains form of tallgrass prairie- Except for the sand dunes and sandhills forms of postclimax tallgrass this ultimate expression of North American 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).

25. 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).

26. 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.

27. 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).

28. 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).

29. 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).

30. 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).

31. 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).

32. 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).

33. 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).

34. 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).

35. 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).

36. 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).

37. 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).

38. 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).

39. 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.

40. 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.

41. 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).

42. 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).

43. 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).

44. 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).

45. 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.

46. Prairie cordgrass- Flowering stalks of prairie cordgrass (famed Chimney Rock, before latest loss of spire height, in background). Morrill County, Nebraska. July.

47. 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).

48. 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).

49. 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).

50. 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).

51. 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

52. 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).

53. 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).

54. 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).

55. 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).

56. 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).

57. 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.

58. 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).

59. 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).

60. 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).

61. 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).

62. 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.

63. 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.

64. 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).

65. 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).

66. 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 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).

67. 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).

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.

68. 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).

69. 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).

70. 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).

71. 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).

72. 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).

73. 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).

74. 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).

75. 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).

76. 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).

77. "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.

78. 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).

79. Swamp milkweed (Asclepias incarnata)- Ottawa County, Oklahoma. August.

80. Shoot apex of foxglove beard-tongue (Penstomenon digitalis)- Ottawa County, Oklahoma. May.

81. 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).

Dominant, Prominent, Major, and Otherwise Interesting Range Plants

of the Tallgrass Prairie

Grasses of Tallgrass Prairie

82. 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.

83. 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.

84. 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.

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

86. 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).

87. 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.

88. 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.

89. 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.

90. 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.

91. 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.

92. 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.

93. 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.

94. 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).

95. 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. Ottawa County, Oklahoma. Estival aspect, June.

96. 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.

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

98. 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.

99. 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.

100. 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.

101. 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.

102.. 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.

103. 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.

104. 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.)

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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105. 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.

106. Next genetic generation of Indangrass- 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.

107. 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.

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

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

110. Switchgrass- Specimen of a bottomland ecotype along the banks of the Bosque River.in north central Texas (Erath County, September).

111. 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.

84. Spikelets on panicle of switchgrasss- Erath County, Texas. October.

112. 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.

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

114.  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.

115. 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.

116. 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.

117. 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).

118. 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.

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

120. 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.

121. 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.

122. 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.

123. 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.

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

125. 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.

126. 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.

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

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

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.

129. 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.

130. 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.

131. 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.

132. 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.

133. 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.

134. 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.

135. 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.)

136. 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.

137. 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.

138. 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).

139. 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

140. 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.

141. 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.

142. 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.

143. 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.

144. 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.

145. 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.

146. 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".

147. 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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148. 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.

149. 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.

150. 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.

151. 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.

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

153. 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.

154. 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

155. 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.

156. 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.

157. Fox 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.

158. 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. Ottawa County, Oklahoma. June.

159. 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

160. 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.

161. 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.

162. 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.

163. 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.

164. 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).

165. 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.

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166. 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.

167. 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

168. 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 such as ditches, lakes, sewage lagoons, and farm and ranch ponds such as the one shown here on Kelly or Lone Oak Prairie, Craig County, Oklahoma. June.

169. Inflorescence of common cattail- Cattail gets its common name from its prominent flower cluster composed of distinctly separate groups of staminate and pistillate flowers. The upper structure (seen here as the gray, curved portion) of the inflorescence consist of male flowers and is called the "cattail"; the lower structure (the brown, cylindrical part) of the inflorescence is composed of female flowers known as the "cat".

Forbs of Tallgrass Prairie

170. Colony of broadleafed (broad-leaf) arrowhead (Sagittaria latifolia)- A wetland or, on some habitats, an aquatic plant that is widespread on wet prairie. It is another monocotyledon. Cherokee County, Kansas. July.

171. Broadleaf arrowhead on tallgrass prairie- Details of leaves and inflorescence on broadleaf arrowhead. Due to presence of a conspicuous corolla the arrowhead species have traditionally been interpreted as forbs, but given that they are monocots a case could be made for regarding them as grasslike plants. Slough along Indian Creek, Newton County, Missouri. August.

172. Early or spring lady's tresses (Sprianthes vernalis) on a badly overgrazed tallgrass bluestem prairie- It would appear that this little orchid is particularily palatable. According to most taxonomic accounts the Orchidaceae is the largest family inthe Angiospermae (flowering plants). Most of these orchid species are in tropical forests (forest generally for that matter) some are grassland species. Two of the more widespread orchid species on tallgrass prairie were included here to provide coverage of more grassland plant taxa. Besides, no other group (family) of "wild flower" or forb can rival the Orchidaceae for sheer diversity of beauty. Ottawa County, Oklahoma. June.

173. Inflorescence of spring or early land's tresses- This dainty orchid inflorescence was on an individual that was a neighbor to the specimen presented in the preceding slide. No other taxon of plants so captures the fancy of wild flower followers than the orchids. None surpass their striking diversity and beauty. The crossover crooner Eddie Arnold sang of sending red roses to a blue lady, but (rose lovers take note) "... if that does the trick I'll hurry back and pick your best white orchid for her wedding gown". Ottawa County, Oklahoma. June.

174. Inflorescence of nodding lady's tresses (Spiranthes cernua)- Some like it early; some like it late. This species of Spiranthes is best distinguished from the preceding S. vernalis while on the range simply by knowing spring from autumn. Ottawa County, Oklahoma. Nodding lady's tresses is a late bloomer while spring lady's tresses comes of age at an earlier season. This specimen was on an old-field that had once been tallgrass prairie. Ottawa County, Oklahoma. November.

175. Beautiful false dragon's-head (Physostegia pulchella)- Yes, that was the long common name listed for this species by the definitive manual for the area (Diggs et al. 1999, p. 770) which called this "the most common Phyostegia of the Blackland Prairie". Another common name for many of the Physostegia species is obedient-plant which originated from the feature whereby individual flowers remain in the position in which they were placed by human hands (or other forces like wind). Physostegia species are members of the mint family (Labiatae) which is often one of the families best represented on prairies and adjoining forests and savannas. This dense stand (which was grown under cultivation for ornamental purposess) illustrated the rhizomatous characteristic of this genus.

176. Inflorescence of obedient-plant (P. virginiana)- This is another Physostegia species found on the Blackland Prairie (Diggs et al., 1999, p. 770). Like the preceding species this one was grown under cultivation.

The Leguminosae is second only to the Compositae in total number of species on North American ranges. A much higher proportion of legumes than composites are valuable forage plants. The nodulated or papilionaceous legumes (generally only members of the subfamily Papilionoideae host nitrogen-fixing Rhizobium bacteria) are the more nutritious and palatable Leguminosae species. With rare exceptions nodulated legumes are forbs and not shrubs. Legumes are higher in energy and nutrients like protein and minerals (especially calcium)  than grasses and most other forbs. Native papilionaceous legumes are desirable members of range plant communities and are often important indicator plants. They are generally decreaser species in which cases their presence on range indicates proper grazing management and higher range condition class. The  extent to which the legume-nitrogen-fixing bacteria symbiosis provides soil nitrogen for other range plants like the grasses has not been determined, but undoubtedly it is of some key role.

177. Indigo bush- Shoot apex with leaves and inflorescence. Nowata County, Oklahoma, May.

178. Inflorescence of purple prairie clover (Petalostemon purpureum)- Several Petalostemon species occur on tallgrass prairie and the ecotone (a transition zone) formed where the borders of the tallgrass prairie and oak-hickory forest merge. The prairie clovers are papilionaceous legumes with the individual five-petal corolla of each flower arranged on a column the entire structure of which is designated a head or, sometimes, a spike. Prairie clovers are quite palatable and relatively sparse compared to the greater relative abundance and herbage production of grasses. The Petalostemon species thus often serve as indicator plants. Prairie clover is viewed as an ice cream species on some range sites where it is able to persist only under light or no more than moderate grazing.

179. Blue wild indigo or blue false indigo (Baptisia australis var. minor)- There are three  species of Baptisia native to the grasslands of the North American Great Plains and Central Lowlands. Blue wild indigo is one of these. It is represented here by a robust  specimen blooming in early morning light on an old horse trap in the Cherokee Lowlands physiographic unit or cherokee Prairie region in southeastern Kansas (Labette County). May.

180. White wild indigo (Baptisia leucantha)- This is another wild or false indigo indigenous to the tallgrass prairies. Like most native legumes it occurs as scattered individuals in dense grassland swards, including the sod of introduced grasses like that of tall fescue (Festuca arundinacea) seen here. Newton County, Missouri. June.

181. White wild indigo- These leaves and flowers are on a specimen growing over 6 ½ feet tall on a Chert Savanna range site of big bluestem and black oak in the Ozark Plateau. Newton County, Missouri. June.

182. Slender bush clover or slender lespedeza (Lespedeza virginica)- This is one of several Lespedeza species native to the central and southern portions of the Prairie Peninsula and tallgrass prairie region. This large legume with its delicate leaves and inflorescences is extremely palatable and generally found only under light grazing and the outside of a fence row. Ottawa County, Oklahoma. September.

183. Wooly or purple locoweed (Astragalus mollisssimus)- Not all papilionaceous, herbaceous legumes are "good guys".  This showy character looks good enough to eat, and so it is thereby creating a poisonous plant problem. Wooly loco causes the classic loco poisoning. The poisonous principle (= toxin) is an indolizidine alkaloid (Cheeke and Shull, 1985, p. 200). This is one of the most costly and spectacular forms of range livestock toxicity. Readers are referred to the timeless work of Kingsbury (1964, ps. 306-311). Remarkably little is known of this historic form of range livestock poisoning as evident by its near absence from the standard veterinary manuals. Wooly loco is widely distributed ranging from Texas to Wyoming eastward to the Dakotas and Kansas. Rocky Mountain National Park, Colorado. Early August.

184. Illinois bundleflower (Desmanthus illinoensis)- Shoots of Illinois bundleflower on a prairie remnant in western Springfield Plateau of Ozark Mountains. This plant was at peak bloom, but in this species blooming is indeterminate and progresses from lower portions of shoot to shoot apex. While Illinois bundleflower is not a papilionaceous legume (it is in subfamily Mimosoideae not Papilionoideae) in some respects this species represents tallgrass prairie legumes as much as any other member of the Leguminosae. Illinois bundleflower is clearly one of the most important native herbaceous legumes in tallgrass prairie across much of the Central Lowlands and eastern Great Plains. The western extreme of this species range extends to Alaska while it is found naturally as east as the Great Lakes Region. Its palatability, large size, wide geographic distribution, drought tolerance, distinctive features, high seed yield, relative ease of propagation, regrowth potential, and competitiveness have made Illinois bundleflower a major species for range reseeding and prairie restoration projects. Soil (Natural Resources) Conservation Service plant materials centers have developed and released numerous accessions of this species over the decades of their diligent work. For descriptions of Illinois bundleflower from perspective of range management and grassland imporvement readers were referred to such basic works as Phillips Petroleum Company (1963, p. 82) or Nicholson (2006, p. 82), Hatch and Pluhar, 1993, ps. 192-193), and Tyrl et al.(2002, ps. 180-181). Ottawa County, Oklahoma. July (early summer); full bloom stage.

185..Beauty and utility- Compund leaves and the infloresence of Illinois bundleflower. These are bipinately compound leaves. The flowers of the Mimosoideae are interpreted as heads of several small flowers each with several stamen. In the Mimoseae tribe of which Desmanthus is a member there are as up to twice as many stamen as petals (Smith, 1977, p. 152). These two slides presented details of floral and foliage features on one of the shoots presented in the two preceding photographs. Ottawa County, Oklahoma. July (early summer); full bloom stage.

186. Young but on their way- Immature cluster or "bundle" of new legumes on one of the shoots of Illinois bundleflower introduced above. It is actually the "bundle" of fruit not flowers that give this important range legume its common name.The dried stigma and stamen are still visible in this cluster of legumes. Ottawa County, Oklahoma. July (early summer); early fruit stage.

187. Nearly there- Clusters of legumes of Illinois bundleflower at midway stage and approaching ripe stage. This plant was a neighbor of the blooming specimen shown in previous slides of this species. All these plants were on a tallgrass prairie remnant in the western part of the Ozark Highlands (Springfield Plateau) in what was formerly a portion of the Prairie Peninsula. Ottawa County, Oklahoma. July (early summer); approaching fruit-ripe phenological stage.

188. There- Photograph of a section of a plant of Illinois bundleflower at fruit-ripe (seed-shatter) phenological stage followed by a photograph of a ripe dehising legumes on this same plant. This particular plant was one of several that were thriving in a field of bermudagrass (Cynodon dactylon) on the flood plain of the Red River. More on this aspect in the next slide ... Cooke County, Texas. July (early summer); peak standing crop and fruit-ripe to seed-shatter stages.

189. Several stories (well, two good ones anyway)- Two plants of Illinois bundleflower in a field of bermudagrass on floodplain of Red River.The competitiveness and relatively large size of Illinois bundleflower was noted previously in this section. Both of those ecologically and economically important characteristics were shown in this photograph of two plants of Desmanthus illinoensis thriving in a dense stand of bermudagrass. Bermudagrass under proper management grows so dense and aggressively that weeds (monocot or dicot) are seldom a major problem with this field crop. The author found it revealing that the only "weed" (defined agronomically as any unwanted plant growing in the intended crop) in this patch of bermudagrass on Red River alluvium was Illinois bundleflower, except for Johnsongrass growing at field edge bordering on the river (background of this photograph). The second story in this slide was a lesson in grazing defoliation from perspective of 1) grazing animal selectivity and 2) morphological/physiological response of plants to defoliation. Close observation of the Illinois bundleflower in this slide revealed not one but two plants. A taller, ungrazed, sexually reproductive plant was growing in front (more forward toward camera) of a shorter, closely grazed (by whitetail deer), sexually immature plant (to rear of tall, fruit-bearing plant). Deer had cropped the one plant of Illinois bundleflower on several feeding bouts as evident from age of scars and production then grazing of regrowth on several partly defoliated shoots. There had been zero feeding on the other (larger, taller, fruit-bearing) plant. This deer-feeding behavior kept the grazed bundleflower in a sexually immature state with more shoots that had less mature stems and leaves (hence, undoubtedly more nutritious forage). Deer converted the grazed bundleflower from a sexually reproductive to an asexually reproductive plant. Both plants survived in their respective states of reproduction and in their morphological forms that were induced by defoliation by deer (or permitted by non-defoliation). Why did deer choose to feed on the one and totally "defer" the other bundleflower that was only inches away from the frequently cropped plant. Was one more nutritious, larger, or of a different plant form at time of first feeding? Did one smell differently? Were any differences due to different genetypes or did a canine (or deer) mark its territory on one bundleflower and not the other? Or was it simply "luck of the draw", a strictly random event? By the way range students, was defoliation or non-defoliation of this palatable prairie forb (herbaceous legume) by free-ranging native ruminants the "natural condition"? Asked in an applied sense, would grazed or ungrazed plants serve as the "control" plant in an experiment (say, for instance, of plant response to drought stress)? Don't jump to conclusions now. Cooke County, Texas. July (early summer); peak standing crop and fruit-ripe to seed-shatter stages.

190. Coming back- Regrowth on an Illinois bundleflower about 16-17 days following total defoliation (no leaves left; stubble height of shoot of six to eight inches) by mechanical shredding (road crew). An unfortunate (?) Illinois bundleflower with a height of slightly less than two feet "got in the way" of a rotary shredder in late June. The bundleflower was stripped of every leaf, young bloom, etc. leaving a spindly, split stalk. Undetured, the brave little bundleflower which, it should be underscored, was rooted in soil moist from recent rains, regrew as shown here in slightly more than two weeks. The shattered basal shoot of this plant died so that there was no living tissue on this above the ground surface. All regrowth was from subterranean tissue. Ottawa County, Oklahoma. July (early summer).

191. Coming back even more- After approximately 35 days following more-or-less total defoliation (except for about a half-foot length of splinttered stalk) by mechanical shredding the never-say-die Illinois bundleflower had regrown densely foliated shoots ass shown in this slide. This and the preceding photograph demonstrated the potential for regrowth (given adequate soil water) of one of the most valuable, native, herbaceous legumes on tallgrass and mixed prairie ranges. Under most systems of proper grazing management Illinois bundleflower would not be completely stripped of aboveground living tissue. On tallgrass prairie meadow that are mowed for hay plants are cut (severed neatly) not twisted off and splintered (ie. beat to death) as with shredding. Hence, injury to mowed plants is much less than damage inflicted by beating and twisting. In other words, regrowth on the Illinois bundleflower shown in these photographs had to make a greater recovery than that which would be required of plants grazed on properly managed ranges or mowed for prairie hay on a commercial basis. Regrowth potential and tolerance to severe (if infrequent) defoliation in Illinois bundleflower makes this an extremely valuable, native prairie legume. Ottawa County, Oklahoma. July (early summer).

192. Another--though littlier--bundle of range joy- Velvet bundleflower (Desmanthus velutinus) on a calcareous soil on Texas' Grand Prairie. This species is considerable smaller (hence less productive) than its "ksssing cousin" Illinois bundldflower. On the other hand, this smaller Desmanthus species is better adapted to less favorable environments (eg. those with shallower, rockier, and generally drier soils). These two Desmanthus species frequently grow in close proximity on the same range as here on this Purves-Dugout complex soil mapping unit of a degraded Rolling Prairie range site (Soil Conservation Service, 1973). Tarleton State University Hunewell Ranch, Erath County, Texas. September. Full-flower phenological stage.

193. Another specimen of velvet bundleflower- Another (of several others) plant of D. velutinus with a more upright form that was growing near the specimen introduced in the preceding photograph presented a more advantageous view of the shoot and inflorescence of this fairly small but palatable prairie legume. Tarleton State University Hunewell Ranch, Erath County, Texas. September. Full-flower phenological stage.

194. Flower and fruit bundled on tallgrass prairie- Two legumes nearing maturity on the apex of a shoot of velvet bundleflower while with an inflorescence lower on the same shoot this specimen looks with anticipation toward even more sexual reproduction. Tarleton State University Hunewell Ranch, Erath County, Texas. September. Full-flower and ripening fruit stage simultaneously on the same plant.

195. Catclaw sensitivebriar (Schrankia uncinata)- This palatable herbaceous legume is in the Mimosoideae subfamily along with numerous “outlaw” woody shrubs that were discussed under the South Texas or Rio Grande Plains savanna type. This decreaser is an indicator species whose presence bespeaks solid stewardship of native tallgrass prairie range. The common name of this palatable and beautiful forb comes from the phenomenon of thigmotropism (= haptotropism), the tropic response of a plant to touch. More precisely it is thigmonasty (nastic movement to touch; nastic movements are those in which the direction of movement is not related to the direction of the stimulus). Whatever it is called the leaflets of the compound leaves fold up under the slightest touch. That condition as viewed here was caused when the plant was inadvertantly brushed 2 feet from the portions pictured. The sensitivebriar in this shot is co-dominant with big bluestem on a Chert Savanna range site in the Ozark Plateau. Ottawa County, Oklahoma. May.

196. Wild senna (Cassia marilandica)- This leguminous forb is a member of the Caesalpinioideae, the smallest legume subfamily in North America. This individual is growing on a bottomland prairie in the Ozark Plateau that is dominated by peaked panicgrass (Panicum anceps). This large specimen is almost a yard tall.

197. Inflorescence and fruit of the wild senna seen immediately above- Ottawa County, Oklahoma. July.

198. Wild alfalfa or scurfy pea (Psoralea tenuiflora)- This is one of the most widely distributed papilionaceous legumes on North American grasslands. It grows from the understory of the western oak-hickory forest of the Ozark Mountains to the Great Plains shortgrass country. Kelly Prairie, Craig County, Oklahoma. June.

199. Flowering shoot of wild alfalfa or scurfpea- Inflorescences of a leguminous forb that is a frequently a local dominant on tallgrass, true, mixed, and shortgrass prairies. This one was in the understorey of an oak-hickory and tallgrass savanna. Ottawa County, Oklahoma. June.

In addition to the leguminous forbs there are also a number of woody legumes--both trees and shrubs--on tallgrass prairie. These woody legumes are in all three subfamilies of the Leguminosae and examples of these were shown below in the section devoted to woody plants. This author elected to organize vascular range plants of tallgrass paririe as to grasses, grasslike plants, forbs, shrubs, and trees with woody species arranged as to families under the more general groups as viewed by rangemen, foresters, and wildlifers. Woody legumes (trees and shrubs of the Leguminosae) were treated below following the Salicaceae.

The Compositae is the largest family of range forbs. Composites are especially plentiful on grasslands. Some are extremely palatable species (decreasers) while others are ecological weeds indicative of disturbances such as overgrazing and drought (invaders).

200. Dense population of compass plant (Silphium laciniatum) at peak bloom- Frisco Railroad right-of-way, Ottawa County, Oklahoma, July.

201. Flowering shoot of compass plant- Typically a decreaser most abundant on properly managed tallgrass ranges.

202. Beautiful colony of Maximillian sunflower on a tallgrass prairie range- This species and compass plant (Heliantheae tribe of Compositae) are decreasers that are so palatable even to Bovidae ruminants that they can usually be found only in ungrazed places like rights-of-way and fence rows or on properly grazed ranges. Wichita Mountains Wildlife Refuge, Commanche County, Oklahoma, October.

203. Inflorescences of Maximillian sunflower- Wichita Mountains Refuge.

204. Engelmann daisy (Engelmannia pinnatifida)- This is another of the composites that is very palatable to grazing animals, including grazers like cattle, that usually prefer grass over forbs. Engelmann daisy has generally been categorized as a decreaser and, though here are other range plant species that experienced observers regard as being more palatable, Engelmann daisy does decline quickly on overgrazed ranges. It's presence on a range indicates that grazing has not been improper very often or very long. Engelmann daisy frequently grows in colonies such as the one shown here, especially in areas like central Texas renowed for it's wild flowers. Engelmann diasy blooms very early in the spring and can arguably be viewed as a cool-season species. Erath County, Texas. (This example was growing in a protected location in the Grand Prairie.) April.

205. Pale purple coneflower (Echinacea pallida)- At edge of a big bluestem-dominated tallgrass prairie in the Prairie Peninsula of southwest Missouri-northeastern Oklahoma this fine specimen of pale purple coneflower graced an equally fine May morning. The Echinacea species are the most common taxon of prairie composites knwon generally as coneflowers. These are also some of the most conspicuous--often striking--forbs on the prairie. Unfortunately for these delightful native wildflowers and those who delight in the beauty they bring to native grasslands the affiliation of Echinacea roots with folk medicine led to gathering of these composites for use in herbal tonics. Root-digging has all too frequently led to local extirpation of local populations of Echinacea species. Any plant species that have valuable, legitimate medicinal properties should be grown as an agronomic or horticultural drug crops using seed as the means of propagation so as not to threaten native populations. Root-diggers of naturally occurring Echinacea "reap where they do not sow" and are a scourge of the human race on par with child-molesters, pimps, and drug-dealers (of which they are a variety). A curse on all your houses! Ottawa County, Oklahoma.

206. Inflorescence of pale purple coneflower - Like most of the leguminous forbs and the last three composite forbs presented, the purple coneflowers are indictor species and generally decreasers. In fact, the best place to find Echinacea species is on well-managed prairie hay meadows and in grassy fence rows where overgrazing by livestock and status-mowing by retirees on riding lawnmowers has not reduced the tallgrass communities to a stomp lots or manicured weed patches of crabgrass. The word "coneflower" is one of the common names that can be misleading (not that scientific names never are!). Echinacea species are the "purple coneflowers" whereas Ratibida species are the "prairie coneflowers" (as was shown below there are other widely accepted common names for some Ratibida species) while Rudbeckia species are often just called "coneflower" (if not something like Black-eyed Susan or Brown-eyed Susan which are names for two separate species). As a general rule, Echinacea species decline under any form of severe defoliation whether by animals or machines and whether too intense, too early, too late, etc. By contrast, many of the Rudbeckia species are less sensitive to abuse and, in fact, some are invaders and indicators of past or present abuse. For example, some Rudbeckia species are often common or even locally dominant on sacrifice areas, old-fields, infrequently traveled lanes or trails whereas Echinacea species are usually restricted to virgin sod (or flower beds of native plant enthusiasts). Newton County, Missouri (on a pristine prairie hay meadow). June.

207. "Coneflower", longheaded coneflower, thimbleflower, or Mexican hat (Ratibida columnifera= R. columnaris)- This multiple common-named, bright prairie denizen even has two ways to spell it's specific epithet! It even has variation in the color of its rays: yellow, red, or purple. These have been used to distinguish between forms, the taxon below variety or the subdivision unit of variety. (Observant viewers should have noted the difference between the rays of the two examples shown here.) "We won't even go there" as this was already too confusing for such a "purty little thang". Mexican hat has customarily been treated as a decreaser or increaser. It is somewhat sensitive to heavy defoliation (typically reaching shorter stature after the first clipping), but it is much too common on improperly managed grasslands (eg. overmowed highway rights-of-way) to be the sensitive indicator plant that Echinacea or Engelmannia species are.

If God had the "inordinate fondness" for composites that He allegedly had for beetles (Coleoptera, largest order of insects) the goldenrods (Solidago species) must have been one of His favorite group of forbs. The goldenrods have to be the botanical equivalent of Darwin's finches or Lick's tits. Flora for states having considerable acreage of prairie, at least before they became the Corn and Soybean Belt, (eg. Illinois, Missouri) often listed 25-30 species of Solidago. Of course, Solidago is a taxonomic nightmare to anyone other than plant taxonomists who "groove out" on such things. Successional response and status of the Solidago species vary, as might be expected from a genus of such diversity. Some are quite obviously rank weeds adapted to disturbed habitats such as old-fields, but others generally have been observed to be restricted to prairies or forest glades in pristine condition. Four Solidago species commonly found on tallgrass prairie were included below just to give students a taste of plant speciation of the range. This short selection began with an overall treatment on one species from organizational level (and photographic scale) of stand, population, or colony through individual plant to that of shoot details of individual plants..

208. Local population of a single goldenrod species- Stand or colony of willow goldenrod (Solidgo salicina= S. ludoviciana= S. patula) on the Grand Prairie in northcentral Texas. This is an example of one of the Solidago species that the experts cannot agree on as to scientific name (or maybe much of anything else). All who view this large group can agree that it is showy (rather attractive to say the least). Some Solidago species grow in colonies or large, local stands such as the one shown here. Other goldenrod species occur more commonly as scattered individuals or in smaller groups than this one. The stand shown here was growing on an old field ("go-back land") on which various native though noxious shrubs had invaded. Willow goldenrod would be either invader (ie. weed) or, perhaps, increaser. At least something was growing to protect what would otherwise be mostly bare soil exposed to ravages of accelerated erosion. Bosque County, Texas. Mid-October; peak full bloom stage.

209. Structure at its edge- Structure of range vegetation is not limited to the scale of plant communities such as range cover types. Even populations of a single species can have a structure consisting of individual plants (or shoots of a clonal species) of various age and size classes, different morphologies, etc. An example of that condition was presented in this view of an edge of the population of willow goldenrod introduced in the preceding two photographs. Whether the smaller shoots growing at the edge were younger, genetically unique plants (or perhaps genotypes of smaller size) or offshoots (clonal shoots) of older, established plants was unknown. Bosque County, Texas. Mid-October; peak full bloom stage.

210. Top-heavy- A single plant with two shoots (or maybe two genetic plants each with a single shoot) of willow goldenrod growing in the colony or population introduced above on an old field on the Grand Prairie in northcentral Texas. The upper portions of these two shoots was so laden with leaves and flower clusters in relation to lower stems that they were bending over in a decumbent form. (Maybe they were of the "weeping willow" race.) Bosque County, Texas. Mid-October; peak full bloom stage.

211. Details, details- Detailed views of two robust individuals of willow goldenrod (Solidgo salicina= S. ludoviciana= S. patula) along with some "sidekicks" of the same species growing on "go-back land "on the Grand Prairie in northcentral Texas. Features of leaves and inflorescencs were clearly visible, but identifiction depended on dichotomous characteristics of stems (Diggs et al., 1999, ps. 408-412 passim) or whichever other authorities were consulted as, for example, Correll and Johnston (1979, ps. 1585-1588 passim). Incidentially, the disparities within Solidgo between these two authoritative sources published 20 years apart was astounding.Plant Taxonomy is about as dynamic as vegetation. Bosque County, Texas. Mid-October; peak full bloom stage.

212. Needleleaf goldenrod (S. gymnospermoides)- The pyramidal inflorescence and narrow lanceolate leaves made identification of this "happy camper" on a prairie hay meadow a "picture book" case. Ottawa County, Oklahoma (on the western edge of the Ozark Plateau). July.

213. Giant or late goldenrod (S. gigantea)- Flowering shoot apex of late goldenrod on a deep sand habitat in the Texas Cross Timbers. As suggested by the specific epithet this specimen grew as tall as the sumac it shared the fencerow with. Erath County, Texas. September.

214. Tall goldenrod (S. altissima)- One of the more common goldenrods in mesic tallgrass prairies and glades of the oak-hickory forest of the Ozark Plateau and adjacent Cherokee Prairie. Tall goldenrod lives up to it's name. This nice colony ranged from three and a half to over five feet in height and gayly proclaimed the onset of autum. Tallgrass prairie in Newton County, Missouri. September.

215. Just what the grasslands needed: another species of snakeweed (and yellow composite)- Annual or prairie broomweed (G. draculoides). This is one of the most common and widespread composites on both tallgrass and mixed prairies, especially in more southernly locations. The two plants (one entire plant plus half the crown of its range mate) presented here was growing on the Grand Prairie of northcentral Texas, a little bluestem-dominated tallgrass prairie community. Annual broomweed is considerably less widely distributed across the Central Lowlands and Great Plains than is broom snakeweed (McGregor et al.,1977, p. 373), but its species range is more than large enough for rangemen preferring grass over annual, weedy composites. Annual broomweed has a far-flung distribution across Texas which has some of the greatest remaining acreage of mixed prairie range, albeit mostly in Poor and Fair range condition classes. Annual or prairie broomweed is an early seral stage colonizer (sometimes a pioneer species) of abused land (eg. overgrazed ranges and those recovering from recent drought, go-back land, highway rights-of-way). Thus this annual broomweed is both an indicator plant as well as a protective (as against soil erosion) and facilitative species that enhances plant succession. Erath County, Texas. October; full-bloom stage.

216. More yeller flowers- Closer-in views revealed an "ocean" of yellow flowers in part of one annual broomweed. Multiply this fraction of one plant times millions of other plants and it takes no imagination to understand why abused ranges--including those just recovering from extreme drought--look more like rape and canola fields than prairies. G. draculoides is in the Astereae (aster tribe) of the Compositae. Erath County, Texas. October;

217. Heath aster (Aster ericoides= Symphyotrichum ericoides)- This palatable perennial commposite is usually the latest-blooming of all the asters with autumn being the season at which dense clusters of inflorescences bedeck many a tallgrass and mixed prairie range. Response of heath aster to grazing is apparently more site-and season-specific than many other range plants. As a rule-of-thumb heath aster is an increaser but on some range sites it responds as a decreaser. Heath aster is palatable, even to grass grazers like cattle, when plants are young (sometimes even up to pre-blooming stage) and become unpalatable at maturity (Phillips Pertroleum Company, 1963). If ranges are grazed when plants of this forb are immature the species responds as a decreaser whereas it responds as an increaser when ranges are grazed later in summer or autumn when heath aster is mature (especially when blooming). Heath aster appears on "go-back land" or "old-fields" relatively early in secondary plant succession, but it persist into the climax (Phillips Petroleum Company, 1963). As such, heath aster is not a good indicator plant for determing seral stages. The plants shown in this and the next two slides were on a degraded Grand Prairie range at a location historically overgrazed by beef cattle. Tarleton State University Hunewell Ranch, Erath County, Texas. October, and obviously full-bloom phenological stage.

218. Blooming aster- Details of inflorescences of heath aster. Plants were growing on a deteriorated tallgrass prairie range in Texas' Grand Prairie. Tarleton State University Hunewell Ranch, Erath County, Texas. October, and obviously full-bloom stage of phenological development.

219. Pink blazing star or pink gayfather (Liatris elegans)- There are a number of Liatris species (though nothing like Solidago). Florae of Missouri and Texas listed nine and twelve species, respectively. L. elegans, shown growing here on a depleted range in the Grand Prairie of northcentral Texas, is a showy species that does well on overgrazed prairies where it's neighbors are other weeds. On this abused range that should support Indiangrass, little bluestem, and big bluestem associated species were curlycup gumweed (Grindelia squarrosa), tumble windmillgrass, and the alien King Ranch bluestem (Andropogon ischaemum= Bothriochloa ischaemum), invaders all. The brillance of the gayfeather inflorescence belies the fact that these species are generally nvaders. Tarleton State University Hunewell Ranch, Erath County, Texas. September.

220. Prairie blazing star or prairie gayfeather (L. pycnostachya)- A dense stand of prairie gayfeather on a prairie hay meadow that was routinely (as in every year) mowed too late (September instead of June). Bluestem prairies (those dominated by the Four Horsemen of the Prairie species) managed as hay meadows in the greater prairie region of western Missouri and eastern to central Oklahoma and Kansas should be mowed from mid June to mid July (southern to northern limits) to keep (or to get) and maintain the vegetation in climax condition. Prairie hay meadows that are dominated by the Four Horsemen will consistently produce maximum yields of nutrients and energy from the land. Under such a "calendar of cutting" the combination of herbage yield and nutritive value of this herbage will be such as to produce the optimum yield of hay for horses and beef cattle, the species for which such hay is outstanding. The most detrimental time for heavy defoliation (as in mowing for hay) of warm-season prairie grasses, especially the panicoid Four Horsemen, is late summer or fall. Harvest of a high proportion of biomass (most of the shoot) of these species at this time (typically when these species are in advanced phenological stages from flowering to seed-shatter) deprives them of adequate time and other resources to regrow photosynthetic tissues and relinish root reserves, grow rhizomes or rootcrowns, etc. that will be essential for growth the next spring. Too-late mowing eventually leads to loss of the bluestems, Indiangrass, switchgrass, and even eragrostoid species like prairie dropseed (this is expressed as winter-killing) and their replacement by increaser and/or invader species such as the healthy colony of prairie gayfeather seen here. The lazy and ignorant farmer who owned this meadow always mowed it (located in northeastern Oklahoma) in September and had weeds for hay. An adjoining meadow was mowed in July and was in "mint-condition" (climax tallgrass prairie). Ottawa County, Oklahoma. July.

221. Indian plantain (Cacalia tuberosa= C. paniculata=C. plantaginea= C. pteranthes)- This is one of the more characteristic herbaceous composites on Flint Hills and Osages Questas tallgrass prairies in higher successional status. This author was not able to find sources that gave response of this species to abusive grazing, but obvious greater abundances of tuberous Indian plantain on well-managed hay meadows, barrow ditches and fence rows, and lightly grazed ranges was practical proof that it is a decreaser or increaser depending on range site. The two plants presented here were growing on a big bluestem-dominated hay meadow in pristine condition. The whole "outfit" was a jewel. Tuberous Indian plantain is another one of those species for which the experts--after better than a century of taxonomic study and publication--cannot agree on its scientific name, even as to genus. Woodson County, Kansas. Late June; late-bloom stage.

222. Leafy base- Basal shoot including lower leaves of Indian plantain. This rosette-like arrangement of basal leaves and the red to brown lines on the stem are very distinctive of this species. Woodson County, Kansas. Late June.

223. Corymb of achenes- The corymb type inflorescence is defined was defined by Smith (1977, p. 292) as a flat-topped or rounded inflorescence in which the pedicels are of varying length". Achene is the most common fruit type of the Compositae. Greenwood County, Kansas (first photograph); Woodson County, Kansas (second photograph). Late June.

224. Inflorescence of Baldwin ironweed (Vernonia baldwini)- This composite (Vernonieae tribe) is a major invader that serves as a common indicator species member of overgrazing on tallgrass prairies and tallgrass-oak-hickory savannas. Ottawa County, Oklahoma. July.

225. White snakeroot (Eupatorium rugosum)- This composite (Eupatorieae tribe) is one of the few poisonous plants of the tallgrass and true prairie range types. White snakeroot was the cause of the dreaded “milk sickness” on the prairie frontiers. Frontiersmen and early settlers were affected (often died) when they drank milk from cows which passed the toxic compounds tremetol or trementone on in lactation. "Milk sick" was a deterrent to settlement of the prairies which were eventually destroyed to utilize their rich brunizem and chernozem soils that became the basis of the Corn-Soybean and Livestock Belt. Ottawa County, Oklahoma. August.

226. Inflorescence of white snakeroot- A flower cluster on the preceding plant. Ottawa County, Oklahoma. August..

227. Western ragweed (Ambrosia psilostachya)- This is one of several species of Ambrosia native to the central grasslands of North America. All of these increase under disturbance (ie. are invader species) with some being large annuals that pioneer denuded spots while others such as western ragweed are perennials that often persist in trace amounts in climax vegetation and become prominent only with disturbances like drought and fire. The fruits (achenes) of Ambrosia species are often major feed sources for upland game and song birds like bobwhite quail. Ragweed pollen is also one of the major allergens (causes of allergy) and sources of hay fever over the vast area called home by these weedy composites. Shoot and/or sneeze. Erath County, Texas. September.

228. Shoot apex of western ragweed- Leaves and immature flower cluster of the most widespread Ambrosia species in interior North American. Western ragweed is one of the most abundant herbaceous invaders on overgrazed grasslands extending from the eastern tallgrass to central mixed prairies. Cross Timbers National Grassland, Wise County, Texas. September.

229. Western ragweed thriving on an overgrazed burn- A wild fire burnt off this portion of tallgrass prairie and these buffalo made the burned area their "base of operation" grazing the palatable big bluestem and Indiangrass as close to the ground as their bovid mouth parts allowed. Simulataneously the "buffers" avoided the foul-smelling western ragweed that was the most common forb. The pehnomenon of grazing selectivity (even by the relatively indiscriminate grazing characteristic of American buffalo) produced this textbook example of localized overgrazing. When the bison "camped out" on the burn they favored the invader species, western ragweed, by improving its ability to compete for resources with the weakened preferred grasses. This demonstrated the reason why sound range management usually requires either 1) adjusting stocking rates to what the burnt acreage can support without overuse (leading in time to overgrazing) or 2) with fixed stocking, firing the entire range so animals cannot overuse the preferred burned portion. Under more-or-less "natural" conditions of free-ranging wildlife and partially burnt-off range overgrazing is unavoidabe. Such combinations of lightening-ignited range fires and native grazing animal behavior occurred repeatedly in the pre-human history of grasslands. This created mosaics of plant communities in various stages of plant succession. It is an example of patch dynamics with resultant maintenance of a maximum (perhaps optimum) diversity of habitats and plant and animal species. In the example shown here, bobwhite quail and various species of passerines (perching songbirds having feet with clasping toes with the first toe pointed backward) will benefit by the increased availability of nutritions achenes produced by the combination of fire and overgrazing by buffalo. Wichita Mountains Wildlife Refuge, Commanche County, Oklahoma. October.

The Asclepiadaceae (milkweed family) is small (based on numbers of species); certainly nothing comparable to the composite and legume families. The milkweeds merit consideration beyond their numbers, however, due to their economic importance and historical prominence as livestock poisoning forbs on range and pasture. In addition, Asclepias species provide one of the best examples of speciation and ecological niches of plants on North Amrerican range. Several Asclepias species were presented below as examples of these two aspects of Range Management and basic Ecology and Evolution. Students should note with interest how many of these species grow on the same range type, in fact on the same range site and often almost side-by-side. The poisonous principle (= toxin or poison) in Asclepias species is associated with two categories of symptoms of poisoning:: 1) gastrointestional-cardiac and 2) neurologic Digestive-cardiac is the opertive form or category responsible for animal poisoning in most of the Asclepias species. Here the asclepiad cardiotoxins are steroids called cardenolides. Toxicity within Asclepias species varies widely widely. This is assumed to be due to different concentrations of cardenolides in the different species (Burrows and Tyrl, 2001, ps. 131-133; see especially Table 12.1). Students were routed to this extraordinary encyclopedia of North American poisonous plants as well as the dated but ever-the-classic Kingsbury (1964, ps. 267-270). The Asclepias inflorescnece is a unique, highly derived structure (set of structures) distinguished by presence of a corona extending beyond (outward from) the corolla. Smith (1977, p. 293) described the corona as an extra set of anatomical structures situated between the corolla and the androecium (entire male structure) that has been interpreted as a modified part of the corolla and/or male unit. The corona consist of five floral units each of which has petal-resembling hood with a beak or crest extending outward from inside each hood. Below and within the surrounding corona the male and female organs are joined or fused to form a compound structure known as a gynostegium . The stamen of the gyostegium are also unique annatomical modifications with side-by-side stamen joined by two connectives or translators that fuse to a central body called the corpusculum. Each of these "half-anthers" secretes a waxy mass of pollen termed a pollliuium. Pollinia are inadvertantly detached by pollinating insects and then almost miracleously inserted inside another milkweed flower to facilitate cross-pollination (Smith, 1977, ps. 187-188). The fruit of Asclepias is a follicle: a podlike, one-carpeled, dry, dehiscent fruit that opens along a single suture. Inside the follicle there are many neatly aligned seeds each of which has an attached coma ( tuft) of ong, silky hairs that act as a natural parachute for dispersion of the seed (Smith, 1977, ps. 187-188, 296).

230. Butterfly milkweed (Asclepias tuberosa)- This showy milkweed is one of 16 Asclepias species listed as having some degree of cardiotoxicity in North America (Burrows and Tyrl, 2001, ps. 125-135 passim; see also the ever-popular Kingsbury, 1964). There are dozens of Asclepias species native to North America. These are especially common on the central grasslands where they are one of the most colorful examples of speciation and ecological niche. Butterfly milkweed is one of the more common (and showy) species in the tallgrass prairie and savanna region. The attraction of attractive spicebush swallowtails (Papilio troilus) to the milkweed illustrated the well-deserved common name of this prairie range forb. Ottawa County, Oklahoma. June.

231. Flowering shoot of butterfly milkweed- Leaves and inflorescence of the milkweed known simply as butterfly weed. This species has very low levels of cardenolides so as to be of relatively low toxicity (Table 12.1; Burrows and Tyrl, 2001, p.133). Ottawa County, Oklahoma. June.

232. Clasping milkweed (Asclepias amplexicaulis)- Clasping milkweed is widely distributed throughout eastern North America from some localities in the eastern Great Plains to eastern New Hampshire. This author has found it to be more common and in counties not shown on official USDA maps or in the various published flora. This was rather surprising given easy identification of this species by its broad leaves that attach directly (and conspicuously so) to the stem, hence the common name of clasping milkweed. .A. amplexicaulis has one of the lowest concentrations of cardenolides and no estimate of toxicity was shown in Burrows and Tyrl, 2001, Table 12.1, p. 133). The specimen shown here and in the next two photographs was growing on degraded tallgrass prairie in the western part of the Springfield Plateau of the Ozark Mountain Region. It is also relatively common in tallgrass-oak-hickory savannahs in this same area. Ottawa County; Oklahoma. Late May; peak bloom.

233. Inflorescence of clasping milkweed- This is one of the least showy or eye-catching of the Asclepias species, but the subtle two-toned hoods of the coronaand conspicuous, ruffled leaves that wrap ever so slightly around the stem make this species an enjoyable beauty for wild flower fans and hard-bitten rangemen alike. Plus with its low levels of toxic compounds (glycosidic cardiotoxins) this species has much to recommend it as a native plant for rural landscaping. Ottawa County; Oklahoma. Late May; peak bloom.

234. Antelopehorn milkweed (Asclepias asperula subsp. capricornu)- Asclepias asperula is one of the widely distributed milkweeds in North America. This species has a discontinuous or disjunct to diffuse biological range with populations centered in the eastern Great Plains and again in the vast region of southwestern deserts (Basin and Range physiogrphic province). There are two subspecies of A. asperula which accounts for this distribution pattern. A. asperula subsp. capricornu is the subspecies known as antelopehorn milkweed. It is the prairie taxon. A. asperula has very high cardenolide concentrations with an estimate that consumption of between one and two percent of animal body weight can cause death (Burrows and Tyrl, 2001, Table 12.1, p. 131). Tarleton State University College Farm, Erath County, Texas. Early May; peak bloom.

235. Inflorescence of antelopehorn milkweed- The flower cluster of Asclepias asperula subsp. capricornu is large and prominent so as to be conspicuous on the range. Its large size, low and sprawling form combined with large inflorescences make this species a favorite (and easily obtained) photographic trophy for wildflower enthusiasts. Tarleton State University College Farm, Erath County, Texas. Early May; peak bloom.

236. American cowslip or shooting star (Dodecatheon meadia)- This member of the Primulaceae or primrose family is the most cherished of all prairie forbs except for the orchids by wild flower afficionados. It is rare except on the best managed prairie hay meadows such as the one where this one grew. Ottawa County, Oklahoma. April.

237. Some mint for flavoring- Wild bergamont (Monarda fistulosa) on a tallgrass prairie savanna in western Ozark Plateau Region (Springfield Plateau section). The mint family (Labiatae) is not one of the larger families of range forbs, but it furnish some colorful and charaacteristic species on most range types including tallgrass prairie. One of the larger and showier of these is wild bergamont, a perennial often described as a subshrub. Typically wild bergamont is not eaten (at least to any appreciable degree) by larger vertebrate animals being instead more conspicuously visited by nectar-seeking insects. Wild bergamont is more common on somewhat disturbed range or, perhaps more precisely, disturbed range that is in various stages of recovery. It is a long-lived perennial and, being a large and showy plant, range folk who routinely walk or ride their favorite prairie haunts find themselves looking for the same plant year after year. Sort of an old botanical pal if you will. Ottawa County, Oklahoma. July, full-bloom stage.

238. Bergamont blooms- Details of the inflorescence of Monarda fistulosa. Each of the tubelike units is a corolla consisting of five fused or united petals. There are at least four subfamilies of Labiatae. Monarda species are in Lamioideae which includes some of the most sought after mints and other flavorings. This does not include oil of bergamont, the defining flavoring of Early Gray tea, which is from a member of the citrus family. Ottawa County, Oklahoma. July, full-bloom stage.

The Umbelliferae is another forb family that is well-represented among range plant communities on tallgrass prairie. Species of this family include both native species and naturalized aliens and range from decreasers of climax vegetation down to pioneer and early colonizing species on old fields and overgrazed ranges. Some examples of the carrot or parsley family were presented in the section below.

239. Poison hemlock, spotted hemlock, or California fern (Conium maculatum)- This intriguing biennial species has the dubious record of being one of the single most toxic plants known. Poison hemlock is enshrined in history as it was an extract from this species that was used to murder the proto-type professor, Socrates. Spotted hemlock is native to Eurasia, but following its introduction in the New World it naturalized across much of North America where it is now a common weed. Poison hemlock is so foul-smelling that it causes little livestock poisoning. Member of the Umbelliferae (parsley or carrot family). The poisonous principal (= toxicant or toxic chemical) is a series of pyridine alkaloids (eg. coniine) which have a mechanism similar to nicotine and can cause death through respiratory failure. This robust (nine feet tall) specimen was growing on California annual grassland in the Russian River Valley. Mendocino County, California. June.

240. Characteristic stem of poison or spotted hemlock- Origin of the "spotted" designation was obvious on this prime specimen. Mendocino County, California. June.

241. Basal stem and tuber of spotted or poison hemlock- These plant parts are useful in identification of this and related members of the Umbelliferae (in contrast to water hemlock for instance). As is the case regarding all poisonous plants in North America readers are referred to Kingsbury (1964, ps. 379-383) and Burrows and Tyler (2002, ps.49-54). Cheeke and Shull (1985, ps. 115-119) is another useful, though less comprehensive, text that was organized around the toxicants rather than by plant families, the traditional approach. This sample was "collected" in the Ozark Plateau, McDonald County, Missouri. June.

242. Water hemlock (Cicuta maculata)- Authorities agree that this is the single most toxic plant in North America. This member of the Umbelliferae is easily and frequently confused with poison or spotted hemlock. Both species have similar habitats and niches (both prefer moist soils; both are usually biennials for example), but water hemlock is native to North America. The poisonous principals (= toxicant) are acetylenic alcohols (eg. cicutoxin and cicutol) with death due to cardiopulmonary arrest. Mechanism is unknown. Again, see Kingsbury (1964, ps. 373-379), Cheeke and Shull (1985, ps. 363-365), and Burrows and Tyrl (2002, ps.54-57 ). Water hemlock was also covered in the Range Plant Handbook (Forest Service, 1940, W52). This colony of water hemlock was growing around a natural spring in the Ozark Plateau. Some of these shoots were over eight feet in height. Ottawa County, Oklahoma. July.

243. Section of basal stem and rootstock of water hemlock- Gross morphology and internal structure of stem and rootstocks have been some of the characteristics used in identification of various members of the Umbelliferae. Ottawa County, Oklahoma. July.

244. Inflorescence of water hemlock- Species of the Umbellifaerae have umbel inflorescences, the feature that was the source for the traditional family name. Umbel is a flat-topped or rounded inflorescence with all flowers borne on pedicels of approximately equal length and arising from a common point on the shoot apex. Ottawa County, Oklahoma. July.

245. Prairie parsnip or prairie parsley, sometimes and more spedifically, Nuttall's prairie parsnip or prairie parsley (Polytaenia nuttallii)- Local stand of a biennial member of the Umbelliferae that is frequently common on southern tallgrass prairies such as those of northcentral Texas. This is a widespread species with a species range from the southeastern Gulf westward to New Mexico and north to the Lower Great Lakes Region and North Dakota, but not making it into Canada. Given the biennial life cycle of this species it could be expected that there are "parsley years" and other years when plants of prairie parsley are "few and far between". The scene shown here was at height of a good year for prairie parsley (came after break of a several-years drought). Although membership in the Umbelliferae (Apiaceae) suggested possibility of toxicity, this species is apparently not a poisonous plant. Polytaenia nuttallii was not included in either Kingsbury (1964) or Burrows and Tyrl (2001). Erath County, Texas. May (mid-spring); full-bloom stage.

246. .Attractive prairie forb- Single plant of Nuttall's prairie parsnip or prairie parsley in the West Cross Timbers part of the Cross Timbers and Prairies vegetational area of Texas. General all-in-one shot of another umbel forb of the central grasslands of North America. Prairie parsley is locally and sporadically a common forb on the tallgrass prairie and Prairie Peninsula Region. Some authors distinguished a Polytaenia texana or P. nuttallii var. texana as distinct from P. nuttallii, but Diggs et al. (1999, ps. 256, 258) lumped all under P. nuttallii. (Texans have a big tendency to see things Texan as unique.) Erath County, Texas. May (mid-spring); full-bloom stage.

247. Ample umbels- Detail of one unit of the umbel inflorescence of prairie parsnip or prairie parsley. Erath County, Texas. May; full-bloom phenological stage..

248. Rattlesnake master, yuccaleaf eryngo, or button snakeroot (Eryngium yuccifolium)- This wierd-looking forb is also a member of the Umbelliferae though of a different subfamily than the two previous characters. Rattlesnake master and button snakeroot were common names given to this plant when it was believed that extracts from its roots were an antidote for snakebite. Yuccaleaf eryngo is a common forb on tallgrass prairies, especially those at higher states of succession like hay meadows. Ottawa County, Oklahoma. June.

249. Inflorescence of yuccaleaf eryngo- Yes, this too is an umbel though in the form a dense headlike structure or a "button" formed by the umbrella pattern of arrangement of individual stalkless flowers. Ottawa County, Oklahoma. June.

A series of photographs of the Eurasian wild carrot or Queen Anne's lace (Daucus carota) growing naturalized on an old field (go-back land) in the Springfield Plateau (western part of Ozark Plateau Region) was presented below as a general example of umbelliferous forbs on tallgrass praire. Bailey (1949, p. 751) described the genus Daucus as consisting of roughly 600 species that were "very widely distributed" and consisting of "more or less weedy plants".

250. Mother Nature's own vegetable garden (with a little help from man)- View of part of an abandoned farm field (an old field or "go-back land") dominated at this season (late spring) by wild carrot, Queen Anne's-lace, or Devil's-plague (Daucus carota). This latter common name was used by Fernald (1950, p. 1104), and it certainly seemed to be the most apt colloquial name for the population of this "pernicious weed" (Fernald, 1950, p. 1104) that had infested this former farmland. Wild carrot is the ancestor of the agronomic (domestic), root crop known as the carrot. Domestication of Daucus carota as an example of artificial selection was treated below in the caption describing the root of this biennial species. This member of the Umbelliferae, the parsley family, is a widespread weed naturalized from Eurasia.. Like almost all annual and biennial species (native and naturalized; grass and forb) wild carrot does best--in fact, probably requires--disturbed ground. It is sometimes a pioneer though more commonly it is a common member of second and/or third seral stages. Wild carrot frequently remains an important species into late seral stages as on the old field shown here that had been taken out of tillage roughly two decades to a quarter century prior to the "photo-plot" shown here. What impact presence of wild carrot has on secondary plant succession through facilitation phenomenon was unknown to this author, but one canno t assume that influence, if any, would be positive or negative. "Weeds" on go-back ground are weeds in name only with forb being the proper or more correct term. Certainly such invading forbs provide some cover and protection of soil against erosion as well as adding soil organic matter. Ottawa County, Oklahoma. Mid-June.

251. Shoots of wild carrot- Specimens of wild carrot emphasizing features of shoot, including leaves and distant view inflorescences (umbels). Fernald (1950, p. 1104) described leaves of Daucus species as "pinnately decompound". The second photograph was of only one abundantly blooming plant. Students should retain this image in their minds (or refer back to this slide) when clusters of wild carrot were presented and described below. Ottawa County, Oklahoma. June; full-bloom phenological stage, peak standing crop..

252. Beauty and study in a blooming umbel- The umbel of wild carrot (most members of the Unbelliferae for that matter) are the ultimate example of a compound arrangement of flowers with the entire inflorescence being divided into successively smaller units or subdivisions of umbels. Bailey (1949, p. 751) described the Daucus inflorescences as "compound involucrate umbels". The general umbel inflorescence type is one having a rounded or flat-topped indeterminate arrangement of flowers all of which are borne on pedicels of more-or-less equal length and arising from a common point on the peduncle. The flower cluster (and arrangement or pattern of flowers) of most members of the Umbelliferae go a degree or arrangement of complexity beyond this ahd have a compound umbel, " a flat-topped or rounded inflorescence in which the peduncles (now called rays) of simple umbels (now called umbellets) are inserted at the apex of the peduncle" (Smith, 1977, ps. 63, 311). In simplier--though not necessarily easier understood--terms, "an umbel of many umbels" (umbels composed of umbels composed of umbels). An umbellet and its supporting ray was shown below. This floral arrangement or pattern of flower organization was shown at progressively finner photographic scale in this series of three slides with the unit organization ending with the individual flower in the third or final photograph. The actual individual (singular) flower of Daucus carota has four or five petals. Ottawa County, Oklahoma. June; full-bloom stage of phenology.

253. Plant-built bird nest- Umbel of wild carrot with ripening fruit. The rays (peduncles) of the umbellets (smallest umbel units) have turned up and inward (toward center of the entire infloresceance, the overall compound umbel) resulting in the "bird nest" physical form of the maturing (and drying) and mature (mostly dried out) compound umbel. No, this photographer never saw a bird build her next in this outfit, but it is this tightly rolled form (most do not progress to this degree of curvature) of the compound unbel that possesses the strong scent of freshly cut carrots. The side view of the compound umbel (second slide) presented the peduncle and showed how the major (largest or first-order) pedicels arise from a common point of origin on top of the peduncle. The first order pedicels (major branches) and second-order pedicels (the rays) of the compound umbel were presented in the first slide. All-in-all, a nifty arrangement; would make a neat pattern on a necktie. Ottawa County, Oklahoma. June; fruit-ripening (though still green) stage..

254. Study of a fruit-bearing umbel- Mature compound umbel of wild carrot of the "bird nest" form. Pedicels and rays (see last two captions above for terminology) curved upward and inward forming a deeply cupped (tightly wrapped) "umbels of umbels". Second slide showed an umbellet and its supporting ray. Those with interest in Weed Science should take careful note of the number of fruits in this one umbellet. Then note how many umbellets are in the compound umbel (first of these two slides). Next scroll back up above and note how many compound umbels are on a single plant of wild carrot. Finally, scroll up to the first photograph devoted to wild carrot and observe how many plants of wild carrot were in one small area (a "photo-plot") of one old field (one tilled unit of abandoned cropland). One need not be a mathematical whiz to visualize why and how weeds are some of the "humble that inherit the earth". Oh yes, almost forgot to mention that each fruit has two seeds. The fruit is a schizocarp that separates into two one-seeded mericarps which come into physical contact to form a suture known as a commissure. The schizocarp is interpreted as an indehiscent (not opening by or along sutures, lids, or teeth) dry fruit because even though it splits along the commisure into its two one-seeded segments, these remain closed and do not shed their single seed (Smith , 1977, ps. 66, 177, 307). Ottawa County, Oklahoma. June; fruit-ripe stage.

255. Wild carrot shoots at ground level- Basal parts of mature shoot of wild carrot featuring leaf axils and colororation of the finely pubescent stem. This is a quite pretty plant, especially for being such an invasive and noxious weed. That lesson in life should be obvious to the intelligent student (and, take it from a veteran classroom instructor, the dumb asses are not worth worrying over). Ottawa County, Oklahoma. June; full-maturity at flowering and peak standing crop.

256. Wild carrot roots (or the carrot before man got ahold of it)- It is generally accepted that Daucus carota is the ancestrial or parent plant of D. carota var. sativa, the cultivated carrot (Bailey, 1940, p. 752). This is a textbook example of Darwin's artificial selection: "Selection by humans of individual plants or animals from which to breed the next generation, because these individuals posesses the most marked development of the required attributes.Typically the process is repeated in sucessive generations until those attributes are fixed in the descendent offspring. Such artificial selection can result in dramatic changes like those that took place in the domestication of plants from their wild forebears" (Allaby, 1998). In the case of Daucus carota var. sativa the most conspicuous "dramatic change" and the "attribute" of agricultural interest was a greatly enlarged--and edible--root. Artificial selection is the foundation of domestication. The roots of naturalized wild carrot presented here were on typical specimens (at least typical of those growing on old fields) so that viewers could compare size, shape, dimensions, etc. of the trait chosen for artificial selection in the original genetic stock to those grown under domestication and industrial agriculture. Wild carrot is a biennial with the enlarged (even by standards of natural selection) root serving as the source of stored food nutrients (energy, protein, minerals) for sexual reproduction during the second growing season of the sporophytic generation. The roots shown here were dug at peak standing crop and the simultaneous full-bloom stage of phenology so that roots had begun to "draw down" as stored food was rapidly expended for demands of fruit production. Unlike the strong role of asexual reproduction in adaptation of many perennial range plants, sexual reproduction is essential for production of the next generation of plants as well as survival of the species over time by periodic recombination of genetic material. Ottawa County, Oklahoma. June; full-flower phenological stage.

There are several Verbena species, generally known by the common name of vervains, on tallgrass prairie and associated range types including oak-hickory-tallgrass savanna and true prairie. These species of the vervain family (Verbenaceae) offered an opportunity to illustrate the important concepts of niche and speciation. A few examples were shown below for pedagogic purposes and the pleasure of general viewers.

257. Prairie vervain (Verbena bipinnatifica)- This is probably the most common species of vervain on the Grand Prairie of northcentral Texas where it blooms early and blesses even degraded range with striking beauty .This is one of the most widely distributed Verbina species with lattitudinal exrtremes of its biological range extending from northern Canada and Alaska to Mexico. It contributes much color to the central grasslands of North America where it is an early bloomer and a welcome sight growing amid dull, weathered range herbage following a long winter. Tarleton State University College Farm, Erath County, Texas. April; full-bloom stage of phenology.

258. Rose or Canada vervain (Verbena canadensis)- This rather low-growing vervain is one of the more common and widely distributed Verbena species on tallgrass prairie, especially in the Ozark through Flint Hills forms. It is less common than hoaryleaf vervain (coming up next) on degraded ranges being more abundant in the protection afforded by fencerows and hay meadows. This species has a wide longitudinal range extending from Florida to New Mexico. With its bright rose to pink inflorescences rose vervain is a favorite among those who prefer to landscape with native plants. The similar appearing prairie vervain presented immediately above is another favorite prairie forb for native landscaping enthusiasts. Ottawa county, Oklahoma. July; full-bloom, peak standing crop.

259. Hoary or hoaryleaf vervain (Verbena stricta)- Sometimes also called slender vervain this taller-growing Verbena species is a showy weed on abused ranges and pastures though not under heavy grazing by sheep which apparently eat it down under such management. This nice specimen was happily growing on a pasture of common bermudagrass (Cynodon dactylon) being grazed by black baldy replacement heifers who obviously did not find it particularly palatable. Ottawa county, Oklahoma. July; full-bloom, peak standing crop.

260. Shoots and flowers of hoaryleaf vervain- Details of leaves and flowers of hoary vervain, one of the more common Verbena species on range and pasture with less than ideal management (not necessarily overgrazing, however). These specimens were growing on a tame pasture of tall fescue (Festuca arundinacea), common bermudagrass, and white clover (Trifolium repens). Ins spite of a dense sward of fiercely competitive agronomic species this prairie forb had held its own. Actually, tame pasture may benefit hoaryleaf vervain. In repeated--though not quantified--observations ove many decades of on-the-ground experience this author has noted that hoary or slender vervain has greater density and larger size on agronomic than on range pastures. Perhaps this species can thrive more competitively, grow to larger size, and produce more fruit (and leave more offspring) due to fertilization, closer grazing, or local soil disturbance associated with more intensive management of grazing lands. Ottawa county, Oklahoma. July; full-bloom, peak standing crop.

261. White flower or nettle-leaf vervain (Verbina urticifolia)- Plants of V. urticifolia are typically some of the largest, if not the largest, of the Verbina species in the tallgrass prairie, oak-hickory association of the eastern deciduous forest, and the ecotone (savanna) between these two range types. Typically none of the Verbina species are abundant other than locally and even then they do not dominate others, have high density, or comprise high proportions of relative cover as do some forbs such as those that form colonies or extensive dense populations. Ottawa county, Oklahoma. July; full-bloom, peak standing crop.

262. A closer look at white flower or nettle-leaf vervain- A photograph of the Verbina urticifolia plant introduced in the preceding photograph that afforded a more detailed view of the shoot, including leaves and the small-flowered yet coarse infloresecence of this species. Nettle-leaf vervain is a frequent rather rank-growing forb on both tallgrass prairie and adjoining tallgrass-oak-hickory savannah. Plants of white-flower vervain in this immediate vicinity seemed to equally at home under post oak, blackjack oak, and black oak (Quercus velutina) or beside beaked panicgrass and Florida paspalum. By contrast, the other vervains (V. stricta and V. canadensis) in this locality were restricted to prairies. Ottawa county, Oklahoma. July; full-bloom, peak standing crop.

263. Carolina, plains, prairie, blue, or white larkspur (Delphinium carolinianum)- This member of the buttercup family (Ranunculaceae) is one of a number of poisonous larkspurs in North America (Burrows and Tyrl, 2001, ps. 1010, 1012). Delphinium species have traditionally been divided into the three groups of: 1) tall larkspurs (roughly two yards or meters in height), 2) intermediate larkspurs, and 3) low larkspurs (generally one meter or yard or less in height). In conjuction with these arbitrary height criteria there are corresponding differences in "general growth form, the root systems, and the seasonal pattern of growth" (Burrows and Tyrl, 2001, ps. 1009). Carolina larkspurr is an intermediate larkspur. The specimen shown here was an individual of Delphinium carolinianum var. virescens. It was growing in a patch of Lindheimer's or Engelmann's pricklypear (Opuntia lindheimeri= O. engelmannii), as if its own toxin was not protection enough. This strikingly attractive prairie forb is widely distributed in North America, extending from the Great Plains to the Atlantic Coast. Carolina larkspur has the most easterly species range of any of the major toxic Delphinium species in North America (Burrows and Tyrl, 2001, ps. 1012-1014). For discussion of larkspurs as poisonous plants readers were referred to Burrows and Tyrl ( 2001, ps. 1009-1017), the most recent "bible " of poisonous plants in North America, Hart et al. (2003, ps. 82-83), and of course, the classic Kingsbury (1964, ps. 131-140). Historically Delphinium has been the most poisonous genus of range plants to cattle (at least those causing greatest economic loss to cattlegrowers) in North America. D. carolinianum has not been one of the more important (more deadly or costly) larkspurs. Larkspur species farther west are a far greater problem than this species of the central grasslands and eastern forests. Carolina or white (blue) larkspur is the major Delphinium species on tallgrass prairie and was included for that reason. Numerous members of the Ranunculaceae are poisonous. Tarleton State University, College Farm, Erath County, Texas. Early May; full-bloom stage.

264. Basal shoots- Leaves and base of stems of Carolina, white, blue, or prairie larkspur. This is the shoot base of the plant introduced in the preceding photograph. Authorities were quoted above to show that morphological factors such as "general growth form" are important in description, separation, and identification of the various Delphinium species.This species--like most members of Ranunculaceae--are conspicuously showy plants. Many of these taxa are commonly used as ornamentals, including larkspurs. Carolina larkspur obviously as merit as a native landscaping plant. Tarleton State University, College Farm, Erath County, Texas. Early May; full-bloom stage.

265. Flowers on a stick- Inflorescence and individual flowers of Carolina larkspur shown at progressively greater detail. The "lark's spur" of Delphinium species is actually the upper sepal (there are five of these which resemble petals) that is elongated and typically raised. Within this calyx are the upper pair of petals which also form spurs. Delphinium is derived from delphinus or dolphin, an allusion of flower shape to a marine mammal (Fernald, 1950, p. 668) so that the common name and scientific name make reference to different orders of the aninmal kingdom. The flower cluster of Delphinium is a raceme Tarleton State University, College Farm, Erath County, Texas. Early May; full-bloom stage.

The Cruciferae (= Brassicaceae), the cabbage or mustard family, also provides a few species of range forbs on tallgrass prairie. The Cruciferae is a major family for field and horticultural crops, including a few ornamentals as well as vegetable and field crops (particularily for oil-seed among row crops), but most of the more notable range crucifers are alien (from Eurasia) invaders. An example of one of these was presented next.

266. Local stand of bastard cabbage (Rapistrum rugosum)- An example of the density and nearly absolute crown cover in a population of this introduced (from the Mediterranean Region) annual crucifer that is expanding its range in North America. Mature lants were shown here at peak-bloom stage. Bastard cabbage was not a major weed at time of this writing, but appears to be spreading and is a cause for some concern. Shoots and lower leaves of Rapistrum rugosum closely resembles those of the various Brassica species, especially B. nigra and B. oleracea. Erath County, Texas. April, full bloom.

267. Plant of bastard cabbage - General habit and morphological features of Rapistrum rugosum at peak bloom phenological stage. Erath County, Texas. April.

268. Base of black mustard- Shoots of two specimens of bastard cabbage showing leaves and branching pattern of this naturalized Eurasian crucifer. These two plants were growing on a disturbance (a construction) site in northcentral Texas (Erath County) in April.

269. Flowers of bastard cabbage- Numerous flower clusters produced on a single plant of bastard cabbage (first of these two photographs) attested to the incrediable capacity for fruit production of this annual Eurasian forb. Details of the inflorescence were presented in the second of these two slides. This is one of several weedy crucifers that can be used for greens.

270. Plenty of bastards- Branches of bastard cabbage with a "zillion" silicles. Capacity for high rates of sexual reproduction was obvious in this photograph of naturalized bastard cabbage in northcentral Texas (Erath County, Texas). May.

271. Study in silicles- Close-ups of the fruit of bastard cabbage. The general fruit type of Cruciferae or Brassicaceae (mustard family) is the silique which is a dry, dihiscent fruit. The silicle is a silique not more than two to three times longer than thick (Smith, 1977, ps. 66, 130, 307). Erath County, Texas. May; seed-ripe stage.

The Scrophulariaceae, the figwort or snapdragon family, also furnishes a number of forbs on tallgrass prairie. Most of these are quite showy and striking in appearance. On tallgrass prairie ranges many , probably most, members of the snapdragon family grown in early spring when there is less competition for light (and other ressources or production-restricting conditions). Some of the more common and conspicuous representatives of this forb family were included in the next section.

272. Painted purple- Two examples of purple Indian paintbrush (Castilleja purpurea var. purpurea ) on tallgrass prairie in northcentral Texas (Grand Prairie). The various Castilleja species are undoubtedly some of the showiest, most stricking of all forbs on the tallgrass prairie. They are also some of the earliest to bloom which most likely is an adaptation to shade given that these shorter or lower-growing plants would be overshadowed by taller-growing grasses, especially the long-shoot tallgrass species. Castilleja species do not appear to be palatable to livestock or wildlife as one seldom sees evidence of grazing on them by vertebrates. They do add beauty to the prairie. These two specimens were growing on shallow, calcareous soil on a sunny west slope. The second of these two plants was a robust, multi-shooted individual.. Tarleton State University, College Farm, Erath County, Texas. Early May; peak bloom phenological stage.

273. Indian paintbrush (Castilleja coccinea)- This, the preceding two plants of purple Indian paintbrush, and the next two forb species are members of the figwort or snapdragon family (Scrophulariaceae). They are all short-day plants that bloom early in the spring before the dominant tallgrasses (at least the "long-shoot" or "culmed" tallgrass species) grow to overshadow and preclude light from them. Indian paintbrush is often the most common spring forb on tallgrass prairies in climax or advanced seral stages (eg. properly managed prairie hay meadows). These pretty specimens grew in the virgin sod of one of the remaining parcels of Burkhart Prairie in Newton County, Missouri. April.

274. Lousewort (Pedicularis canadensis)- This member of the snapdragon family blooms early in the spring before it is "overtopped" by larger, taller-growing grasses and summer or fall forbs. Lousewort and Indian paintbrush (and many other spring-flowering forbs) impart a distinctive vernal aspect to tallgrass prairies and oak-hickory and tallgrass savannas. Missouri State Prairie Park, Barton County, Missouri. April; full-bloom phenololgical stage.

275. Shoot apex of lousewort at full-flower- Inflorescences and leaves of lousewort growing in virgin sod of tallgrass prairie. This species is indicative of climax or high seral stages in Andropogon-Sorgastrum-Panicum-dominated grasslands in the Central Lowlands physiographic province. Missouri State Prairie Park, Barton County, Missouri. April.

276. Cobaea (often, Cobey's) beardtongue or Cobey's penstemon (Penstemon cobaea)- This member of the snapdragon family is quite palatable to grazing animals. It has traditionally been categorized as a decreaser. It is an indicator species whose presence on a range usually signifies proper use (or somewhat on the light side of proper use). Hunewell Ranch, Tarleton State University, Erath County, Texas. May.

277. Two more specimens- Two more plants of Cobey's beardtongue to show whole plant (shoot) habit and morphological characteristics of this beautiful prairie forb. These fine representatives were growing on a shallow calcareous microsite on a previously overgrazed range on the Grand Prairie of northcentral Texas. (This range currently was not being overgrazed--in fact, it was not being grazed by livestock at all--otherwise these palatable babes of the prairie would not have been there.) Tarleton State University College Farm, Erath County, Texas.April: full-bloom stage.

278. Shoots and flowers of Cobey's beardtongue- Two plants of cobey's beardtongue showing shoot features at closer distance. Second and third photographs were of same plant with third slide a close-up of two opened and two unopened flowers. Note last year's dead shoot to right of this year's shoot. These showy specimens were on shallow calcareous microhabitats on a formerly abused range of northcentral Texas Grand Prairie. Tarleton State University College Farm, Erath County, Texas.April: full-bloom stage.

279. Cutleaf, yellow, or plains evening primrose; halfshrub sundrop; or yellow sundrops (Oenothera serrulata= Calylophus serrulatus)- This warm-season, perennial member of the of the Onagraceae (evening primrose family) is a palatable forb classified as a decreaser, although Tyrl et al. (2002, p. 239) regarded it as a species of mid-sere. The more frequently used common name of halfshrub sundrop was derived from the semi-woody basal shoot (ie. probably best described as a suffrutescent species). This evening primrose species can serve as an indicator species of overgrazing, but is usually not abundant enough to be an important (at least not a major) forage plant. Grand Prairie, Erath County, Texas. April, full-bloom phenological stage.

280. Close-up of one of the characteristic prairie forbs- Details of leaves and inflorescences of the specimen of halfshrub sundrop or cutleaf evening primrose shown in the preceding two photographs. Erath County, Texas. April, full-bloom stage.

281. Fringeleaf false-petunia (Ruellis humilis)- This minor, even incidential, prairie forb was included to represent the Acanthaceae (acanthus family) thereby presenting still yet more botanical diversity of the tallgrass prairie to viewers. This also reinforced the lesson that there are many families of range plants which contribute relatively small numbers of species yet which contribute to the rich flora of this range cover type. Ottawa County, Oklahoma. July.

282. Pair of plantains- Two species of Plantago growing on a disturbed tallgrass paririe range. The taller and all-around larger plant was tallow-weed or red-seeded plantain (P. rhodosperma). In front of the conspicuous plant of red-seeded plantain was a smaller individual of Heller's or cedar plantain (P. helleri). Both species are native annuals, and both indicate degraded range. They are examples of indicator species; in this case, grazing indicators. Tarleton State University College Farm, Erath County, Texas. April.

283. Heller's or cedar plantain (P. helleri)- This is the individual growing directly in front of the red-seded plantain specimen presented immediately above. This is the smaller-sized of these two species. They both hold some soil in place and provide some oxygen while fixing and sequestering carbon atoms. Otherwise, one would be hard put to come in with much practical or economic value to these native weeds that are indicators of deteriorated range. Tarleton State University College Farm, Erath County, Texas. April.

284. Purple passionflower, apricot-vine, or maypops (Passiflora incarnata)- An herbaceous vine growth form, a viney forb on tallgrass prairie, and a member of a small family with a unique (and showy) inflorescence are combined in this member of the Passifloraceae. Leaves of this species are large and three-to five-lobed. Shoots have large tendrils so that this species s an herbaceous climber. Such a combination of plant features made this species an interesting range plant to include in this presentation of prairie forbs. Plants of purple passionflower can be found growing either individually or in groups. This species does not appear to be eaten by livestock or game animals, at least not under light to moderate grazing, although it is more often found in fencerows. Ottawa County, Oklahoma. July; full-bloom stage.

285. A passion for beauty where it is found- Views of the inflorescence of maypops or purple passionflower. The corolla of Passiflora species consist of series of "thread-like filaments" (Smith, 1977, p. 125) arranged in a three series around a flesh-colored crown (Fernald, 1950, p. 1042). All-in-all a most distinctive and attractive inflorescnce. Ottawa County, Oklahoma. July; full-bloom stage.

286. Fruit of the herbaceous vine- Fruit of purple passionflower. Fruit type of the Passiflora genus is a many seeded berry (Fernald, 1950, p. 1042). It is the fruit of this species to which the term maypops is applied. Ottawa County, Oklahoma. September :ripening though still immature fruit stage.

Shrubs and Trees of Tallgrass Prairie

287. Two decreaser shrubs of tallgrass prairies and oak-hickory savannas are redroot (Ceanothus ovatus) and ...

288. New Jersey tea (C. americanus)- New Jersey tea got its name from the practice of brewing a drink from its leaves by backswoodsmen along the Atlanic Coast and up above the Fall Line in the eastern deciduous forest in which this shrub was a dominant understory species. Both plants in Ottawa County, Oklahoma. June.

289. Rootstock of redroot- The large rootstocks of New Jersey tea (and sometimes the smaller rootstocks of redroot seen here)  were called “grubs” by backswoodsmen who had to remove these organs from their fields carved from the virgin forests for certain crops (eg. sweet potato). According to the record left by frontier historians this was origin of the term “grubbing”.

290. A fruitful plant of New Jersey tea and its fruit - Specimen of C. americanus with a heavy fruit crop (first photograph) and detail of a cluster of this fruit (second photograph) growing on a dune in Nebraska Sandhills. Fruit still immature. Cherry County, Nebraska. June.

291. Prairie rose (Rosa setigera)- Shoot of the low-growing, native rose of the tallgrass prairie complete with an exceptionally dense group of the striking flowers of this species. Ottawa County, Oklahoma. May.

292. Sand, western sand, or sandhill cherry (Prunus bessey= P. pumila var besseyi) This member of the Rosaceae calls the Nebraska Sandhills home. It was originally recognized as a separate species and named in honor of the famous botanist Charles E. Bessey, Father of the American Plant Ecology and mentor to such greats as Frederic E. Clements. Today sandhill cherry is regarded as a taxonomic variety of the widely distributed sand cherry which was one of the shrubs on the sand dunes of Lake Michigan where Henry Chandler Cowles was probably the first American ecologist to describe plant succession and recognize it as a dynamic phenomenon. It was from Cowles' use of dynamic that Clements took the concept and made famous his model of "dynamic vegetation". Cowles' major professor was John Colter who wrote famous flora of the Western Range Region including the classic, Botany of Western Texas (Colter, 1891-1894). But back to the range plant ... Sand cherry is one of several low-growing shrubs in the postclimax tallgrass prairie of the Nebraska Sandhills. It is also distributed as far north and east as Newfoundland and Qubec. Western sand cherry undoubtedly provides some browse and nutient-rich fruit (plums are drupes) for wildlife or even livestock. All Prunus species should be considered potentially toxic (amygdalin, a cyanogenic glycoside, that breaks down to hydrocyanic acid is the poisonous principle), but the smaller P. bessey poses less threat than larger species such as P. virginiana and P. serotina. Cherry County, Nebraska. Late June.

293. Big tree on the prairie- Beautiful and massive specimen of bur oak that made its home in the floodplain of the Bosque River in the Western Cross Timbers of northcentral Texas. Bur oak is obviously a minor member--especially when compared to the dominant post and blackjack oaks--of the scattered forest and woodland communities in southern portions of the tallgrass prairie region. In more northernly parts of this vegetational realm bur oaks replace blackjack and even post oak as the dominant oak. In these northern parts of the Prairie Peninsula and tallgrass-true prairie region bur oak would be second--if that--only to eastern cottonwood as a dominant tree both on grassland and isolated forest communities such as gallary forests. Bur oak is the most widely distributed of all the oak species that have the role of hardwood dominant across the North American central prairies (McGregor et al., 1977, ps. 39-41). Post oak and blackjack oak barely extend northward to Iowa whereas bur oak extends to Ontario. Hamilton County, Texas. July.

294. Bur(r) or mossy cup oak (Quercus macrocarpa)- Young leaves and catkins of the dominant oak of the northern tallgrass, true, and mixed prairies northward from central Kansas. Bur oak can be interpreted as the northern ecological equivalent of post and blackjack oaks as   the aboreal dominant of the grassland-deciduous forest ecotone (Vankat, 1979, p. 221).  This species produces the largest acorns of any oak in North America and it is the oak of the famous oak groves and savannas of the northern grasslands (Peattie, 1938). Seedlings rapidly send their tap roots deep into the fertile prairie soils and become quickly established after germination (Weaver, 1968, ps. 135-139). This genetic adaptation to drought combined with the species’ thick fire resistant bark (Allen, 1967, p. 15) make it admirably suited to drought- and fire-prone prairies. It’s range extends far south of it’s region of dominance into central Texas where it sometimes dominates bottomland savannas. Hamilton County, Texas, April.

295. Leaves and fruit of buroak- Burr oak (either one or two "rs" are used) is regarded as having the largest acorn of any Quercus species in North America. Like other white oak species buroak bears fruit every year (versus a two-year cycle) in the red or black oaks. The combination of large acorns and production each year makes this species one of the most valuable producers of mast in the eastern deciduous forest (ie. the eastern part of North America). Burroak is also known as mossy-cup oak, both common names in reference to the conspicuous tapered tips of the scales of the cup which present a fringe-like appearance. The geographic range of. Q. macrocarpa extends from eastern Canada (New Brunswick and Quebec) to Texas. Hamilton County, Texas. September.

296. Tough tree in "hard scramble" country- Small and misshapen (stunted runts) chinquapin (= chinkapin) oaks on the shallow soil of an exposed south in Cherokee Prairie. Chinquapin oak can development into large, even immense, stately trees given its optimum environment. Less than ideal habitat can still make for survival (defined as successful reproduction) as on this marginal site for this species. One of several common names for Quercus muhlenbergii is rock oak. This and the specimens shown in the next slide typify that habitat feature. Throughout most of the Prairie Pninsula chinquapin oak is a major hardwood species in savanna and forest vegetation. In fact, chinquapin oak has a species range throughout the Prairie Peninsula and tallgrass prairie region that is larger than any other dominant oak except for bur oak (Barker et al., 1977, ps. 39-41). Rogers County, Oklahoma.

297. Two oaks on a dry branch of the Modoc- Two adult chinquapin oaks of mid-size on a dry, stoney stream bank in Ozark Highlands. Note in both these individuals as well as those shown above that chinkapin oak frequently branches close to the ground with these oldest, largest limbs persisting throughout much of the trees' lives. This feature is characteristic of trees in general that develop in the open as on prairie or savanna. Ottawa County, Oklahoma. October.

298. Springtime for chinkapin oak- Catkins and young leaves of chinquapinoak as it emerged from dormancy began its annual cycle of life again. Dallas County, Texas. March.

299. Chinkapin oak- Two samples of leaves and fruit of chinquapin oak. First of these two slides was of the "hard scramble" specimen used to introduce this species (Rogers County, Oklahoma. October). Second sample of foliage and acorns was from a well-watered, well-nourished individual ( Dallas County, Texas. October).

300. Ozarks blackjack- Mature specimen of blackjack oak (Quercus marylandica) on a hardwoods-tallgrass savanna in western Springfield Plateau portion of Ozark Mountains. Understorey had been highly modified by overgrazing followed by overmowing. It was revealing that the major grass next to trunk of this tree was western wheatgrass (Agropyron smithii). Autumn coloration. Ottawa County, Oklahoma. October.

301. "Trees on tallgrass prairie? Yes, (at least by Kansas standards)"- Two views of eastern cottonwood (Populus deltoides) growing on tallgrass prairie. Some of the bent, horizontal trunks (most of them were cut off by the Epson printer*) of the second slide were black willow (Salix nigra). These two members of the Salicaceae are often associated with each other on moist habitats in both tallgrass and mixed prairie communities. Saline County, Kansas. November. *In this photographer's experience Epson printers are fast, but a clearly inferior product for scanning slides. This author's Epson Perfection V700 photo printer overexposed many slides and yet failed to show clearly many clear slides such as these two that had proper lighting. Worse yet, the Epson Perfection V700 scanner over-cropped almost all slides. This overpriced equipment is far from "perfection". R. E. Rosiere recommended against purchase and use of Epson scanners.

302. Crowns on the autumn prairie- This is about as colorful as Kansas prairie gets. It may not be the "yellow brick road", but cottonwood leaves on cow paths seemed mighty purty. Eastern cottonwood is the State Tree of Kansas. Butler County, Kansas. Late October.

303. Foliage and fruit of prairie silvan queen- First of these two photographs was young shoots (two small seedlings or small saplings) of eastern cottonwood. They had been flattened by running water from recent spring (June) rains. (They straightened up.) Second photograph was the typical deltoid leaf with ripe fruit ( a capsule) of eastern cottonwood. Cottonwood capsules are described as comose, an adjective referring to having a coma (a tuft of hairs). Cottonwood derived its common name from the phenomenon of conspicuous, usually copious (and inconvenient) shedding of the comose ("cottony") fruit. All members of the Salicaceae are dioecious so that a tree is male or female and thus produces only staminate or only pistillate flowers. Of course only female plants produce fruit so that occurrence of the "messy fluff" comes only from the ladies. (Some range men find girls troublesome while others enjoy the mess.) Female catkins from this same plant were shown in the next photograph. Erath County, Texas. May.

304. Cottonwood "eggs"- Pistillate catkins of eastern cottonwood. These catkins were from the same tree whose leaf and fruit were shown in the immediately preceding photograph. All members of Salicaceae (willow or poplar family) are dioecious so that catkins are unisexual (ie. all catkins from each individual plant [= each genetotype] are either male or female flowers). Erath County, Texas. April.

305. Prairie willow (Salix humilis)- Female plant (in peak fruit shatter) of prairie willow. A prairie in the Ozark Plateau. Ottawa County, Oklahoma. April. This is one of the most widely distributed willow species on central North American prairies.

306. Ain't they sexy!- Unisexual flowers or prairie willow. In was explained above in discussion of eastern cottonwood that species of the Salicaceae (willow or poplar family) are dioecious. The flower cluster of the Salicaceae is a ament or catkin, "an inflorescence characterized by typically unisexual.apetalous [absent petals], bracteate flowers in a pendant cluster (Smith, 1977, p. 291). In the first of these three slides both pistillate or female (left shoot) and staminate or male (right shoot) catkins were present. This was side-by-side occurrence was the natural position of the female willow and the male willow illustrating ready (handy) pollination in Salix humilis. The second slide was of the female shoot and catkins. Third slide was male. These plants were growing at edge of an oak-hickory forest and small prairie in the western Ozark Plateau. Newton County, Missouri. April.

307. Flowering before the tallgrasses grow tall- Fruit (comose capsules) of prairie willow on edge of an oak-hickory forest and a small prairie in the western Ozark Highlands. Newton County, Missouri. April.

It was remarked above that in addition to leguminous forbs there are also a number of woody legumes--both trees and shrubs--on tallgrass prairie. These woody legumes are in all three subfamilies of the Leguminosae ranging, for example, from the generally invasive species like honey mesquite (Prosopis glandulosa) of Mimosoideae on the southern prairies and the more widely distributed honey locust (Gleditsia triacanthos) of the Caesalpinioideae to shurubs like the attractive eastern redbud (Cercis canadensis), also in the Caesalpinioideae, and northern or fragrant mimosa (Mimosa borealis) of the Mimosoideae. Some examples of these woody legumes were presented in the following section (rather than above with herbaceous legumes )so as to arrange vascular prairie species in the more general groups of range plants (ie. grasses, grasslike plants, forbs, shrubs, and trees). Then various species of woody prairie plants were arranged as to families and shown under these general groups.Some woody members of Leguminosae followed.

308. Bright pink and catclaws- Fragrant or northern mimosa (Mimosa borealis) is one of the smaller shrubby legumes on tallgrass prairie. This shrub is rarely abundant occurring instead as widely scattered individual plants. It is also rather restricted in its biological range being primarily a range plant of the southern prairies as, for instance, in tallgrass prairies of northcentral Texas and on mixed praire in the Southern Great Plains . Although bearing a specific epithet that designates it as "northern" Mimosa borealis does not grow north of southern Kansas (McGregor, 1977, p. 154). Likewise, this wispy shrub does not stand out among other species to the extent that larger or more showy woody legumes (eg. eastern redbud as shown below). The two individual plants shown here were in full bloom in a Texas mid-spring. Tarleton State University Hunewell Ranch, Erath County, Texas. May.

309. Dainty and "purty" on the prairie- Two leaders of fragrant mimosa at full bloom and with young spring leaves before the ravages of wind, hail, drought, and browsing take their toll of this small shrub later in its growing season. Erath County, Texas. April; full-bloom phenological stage.

310. Beauty if you look for it- The small and colorful inflorescences of fragrant mimosa. The flowers of each of the three subfamilies of Leguminosae are distinct and different from each other. The flower cluster of the Mimosoideae are arranged as relatively compact heads with numerous, individual, cylinderical flowers each of which has brightly colored stamen extending beyond the petals of its tubular corolla (Smith, 1977, p. 152). Erath County, Texas. May.

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311. Little stuff for a shrub- Small leaves and legumes characterise fragrant mimosa. This shrubby legume does possess the catclaw-shaped prickle of other Mimosa species, but these are less bothersome to those moving around them than is the case for most of the others. First of these two photographs was from a plant in Quay County, New Mexico in June. The second photograph was of a plant in Erath County, Texas in July.

312. Eastern redbud (Cercis canadensis)- This native shrub or small tree of the Caespalpinioideae subfamily of the Leguminosae has a geographic range from Ontario to the mountain ranges of the general Chihuhuan Desert region. Redbud is often an associate of flowering dogwood in the eastern decuiduous forest where the two dominate the second woody (= under shrub) layer. Farther west eastern redbud occurs without flowering dogwood either as a more or less solitary shrub on grasslands or, more commonly, associated with the dominant shrubs of those locales. The deer-browsed individual seen here was growing on the Texas Edwards Plateau where it is a very valuable browse species associated with numerous shrubs like true mountain mahogany, Texas buckeye (Aesculus glabra), Texas persimmon (Diospyros texana), agarito (Berberis trifoliolata), live oak, whitebrush (Aloysia lycioides), and other legumes like honey mesquite (of course) and Texas mountain laurel (Sophora secundiflora). That's just for starters! Kerr Wildlife Management Area, Kerr County, Texas. June.

313. The red "buds" of redbud- The namesake "bud" of this brightly colored native legume is actually the open floral bud with its fully developed inflorescence. Branches of this shrub are to a great degree covered by multitudes of this bright pink or light red flowers which senese before any of the baby cordate leaves appear. The woods from the eastern deciduous forest through to the parts of the Great Plains such as the Edwards Plateau are ablaze in early spring with flowers of redbud, often accompanied by other spring-blooming shrubs such as flowering dogwood (Cornus florida). The example presented here was in the Western Cross Timbers of northcentral Texas (Erath County). March.

314. Bounteous beauty of spring up close- Close-up views of the inflorescence (and individual flowers thereof) of eastern redbud at full bloom. From other than very close examination the flowers of redbud appear to be papilionaceous. They are not in spite of an "almost papioionaceous corolla" (Smith 1977, p. 153). Instead redbud has the flower of the Caesalpinoideae and not the Papilionoideae subfamily of Leguminosae. Erath County, Texas. March.

315. Dull-colored fruit of redbud- It is the fruit type, legume, and not the flower type or structure that distinguishes and designates members of the Leguminosae. These fully ripe and starting-to-dehisce (split open) legumes were on a redbud growing on a clcareous outcropping in the Grand Prairie of northcentral Texas. Tarleton StateUniversity Hunewell Ranch, Erath County, Texas. October; fruit-ripe phenological stage.

Insert redbud photographs from Texas edwards plateau

316. The thorniest tree in North America- Views of trunks of honey locust (Gleditsia triacanthos) showing the extreme degree of thorn development and the instantaneous identification characteristic of this species. Honey locust is arguably the most widely distributed leguminous tree on the tallgrass and true prariries of central North American. Honey locust extends across much of the Central Lowlands physiographic province ranging from Texas to North Dakota (McGregor, 1977, p. 157). Hioney locust grows naturally as far west as the Staked Plains though this is more a matter of biological (species) range than as a meaningful component of grasslands on the Southern High Plains in texas and New Mexico. It is also a forest tree as far east as the Middle Atlantic states like New York (Fernald , 1950, p. 885). Honey locust is of course far less common as a prairie tree than cottonwood, willow, and several of the oaks (post, blackjack, and bur), but it is one of the more abundant trees after these major species. The one big exception to this is when honey locust invades abused (overgrazed, overmowed, underburned) prairies or go-back land. Honey locust commonly invades abandoned farrm ground (old fields) to such degrees of density and cover as to become a major brush species. It is sometimes been referred to as the "mesquite of the Mid West". In addition to the effectiveness with which honey locust spreads to considerable cover on abused or abandoned land as well as the "triple-branched" (sometimes quadrauple branching), gigantic thorns that sometimes grow to dimensions resembling small branches that make honey locust a dreaded brush species in the eastern tallgrass and true prairie region. Honey mesquite does make a desirable shade tree, and the thorns are not frequently shed (though the case could be made that one such is more than enough). It produces wood of beautiful color, distinctive grain, and remarkable hardness. This latter is especially so by standards of legume wood. Honey locust has been widely planted in shelterbelts and as a shade tree in semiarid areas of the Great Plains. G. triacanthos var. inermis is a natural taxonomic form lacking the objectionable thorns that is preferred for shelterbelt and ornamental purposes. Unfortunately from this arborcultural goal, thornlessness is neither a genetically/phenotypically fixed trait nor a simple recessive trait (Michener 1986). Honey locusts that are thornless in their youth will sometimes develop thorns as they mature so that more thorough study resulted in rejection of the taxon G. triacanthos var. inermis (Michener 1986). Personally this author, who grew up around honey locust (including fighting them as brush) and admires them, has found their thorns much less of a nusiance and hazard than typical university administrators, but such a comparison is repulsively unfair-- to the tree. The tree trunks in these two slides were in an ecotone in the western Ozark Plateau between a bottomland forest dominated by western hackberry and American elm within the general oak-hickory forest and a big bluestem-dominated tallgrass prairie that was an island of Cherokee Prairie grassland. Ottawa County, Oklahoma. April.

317. Represntative of the Caesalpinioideae- Leaves and inflorescences of honey locust growing in a savanna between the overall oak-hickory forest of the Ozark Plateau and bluestem prairie of the Cherokee Prairie. Ottawa County, Oklahoma. May.

318. Honeyed flowers up close- Details of flower clusters of honey locust. Close-in views of inflorescences presented in the preceding photograph. The corolla of individual flowers in the Caesalpinioideae somewhat resemble the papilionaceous corolla of the Papilionoideae, but this resemblance is less in honey locust than in some other caesalpinioid species like rebud showed above. Ottawa County, Oklahoma. May.

319. Sexy and thorny- The typically high yield of members of Caesalpinioideae was displayed in these two photographs of legumes of honey locust. The large, flat, and twisted legumes of honey locust render it readily identified in the field-- as if the large, exagerrated thorns were not adequate for this purpose. High yields of many-seeded legumes account for successful invasion of honey locust on overgrazed pastures and ranges as well as old fields. In the authors emperical observation wildlife and livestock make remarkably little use of these fruits as compared, for example, to those of mesquite. Extent, if any, of seed consumpition was unknown to this writer Martin et al. (1950) made no reference to use of honey locust--as either browse or mast--by wildlife. First slide from Ottawa County, Oklahoma in September; second slide from Benton County, Arkansas (along War Eagle Creek) in July.

320. Honeysuckle on the prairie? Yep.- White or bush honeysuckle (Lonicera albiflora) is one of the more distinctive shrubs of the tallgrass prairie vegetation in Texas' Grand Prairie. This mdeium-size shrub typically occurs as widely scattered (wide-spaced dispersion) individual plants not as groups of plants. Nonetheless, bush honeysuckle is a characteristic plant of tallgrass prairie on calcareous soils. In fact, bush honeysuckle is an indicator plant whose presence indicates limestone or chalk parent material. This species appears expecially fond of limestone outcrops, the habitat on which both of the plants presented here were growing. The plant in the first of these photographs had grown up on a dead snag of blueberry or Ashe juniper (Juniperus ashei). The example of foliage and flowers in the second photograph was a portion of a very large plant that was growing out of a narrow crack in a limestone shelf. Tarleton State University Hunewell Ranch, Erath County, Texas. April. Full-bloom phenological stage.

321. A native shrub that makes a perfect ornamental- Details of inflorescences of bush or white honeysuckle growing on tallgrass-dominated, grassland range on the Texas Grand Prairie. These two photographs were of flowers on the plant shown in the immediately preceding slide. It would be impossible to come up with a more stikingly beautiful shrub to adorn a prairiescape than this species (which is to be expected when God was the landscape designer). Native plant buffs in the central prairies of Texas should make much use of this species when landscaping their yards, especially around ranch headquarters. It should also adorn visitor centers and campgrounds of parks and wildlife refuges. Tarleton State University Hunewell Ranch, Erath County, Texas. April. Full-bloom phenological stage

322. Honeyed fruits of the prairie- Tips (terminal ends) of shoots of bush or white honeysuckle showing the distinctive arrangement of fruit (typically four fruits) centered atop two opposite and touching leaves. This arrangement of terminal leaves is a characteristic of several Lonicera species (see line drawings in Vines, 1960, ps. 952-957). Everything about this striking and picturesque shurb from bark to eye-catching flowers to this unique feature of fruit-bearing shoots is distinctive and attractive. The flowers undoubtedly attract a diverse array of pollinators and the fruit is bound to be eaten by various species of wildlife. The most commonly noted use of the fruit is as a folk medicine for emetic and cathartic uses (Vines, 1960, p. 952; Diggs et al., 1999, p. 508). Tarleton State University Hunewell Ranch, Erath County, Texas. September. Immature fruit stage of phenology.

Organization note: Trumpet or coral honeysuckle (L. sempervirens) is another native shrub in the Texas Cross Timbers and Prairies vegetational (land resource) area, but this species grows on sand, especially deeper sandy soils, of the Cross Timbers. Thus these two Lonicera species do not usually occur in close proximity to each other except where Cross Timbers and Grand Prairie vegetation are conterminous in this patchwork of grassland and savanna range vegetation. Trumpet or coral honeysuckle was presented with the Cross Timbers cover type under Tallgrass Savanna in the Grasslands biome heading.

323. As if scratches from sand weren't enough- Western poison ivy (Rhus rydbergii= Toxicodendron rydbergii= T. radicans var rydbergii) in heart of the Nebraska Sandhills. Another species that the "experts" cannot agree on as to its proper scientific name. Those sensitive (allergic) to this species can agree that it gives a bad itch and an almost irresistable urge to scratch. Fortunately this photographer is "immune" to the toxin that causses dermitis to others who enjoy the sandhills. Range animals sometimes browse this species and obviously birds make use of the fruit. Two-legged range folk can enjoy this sight-pleasing small shrub of the sand dunes, especially from safe distance afforded by the worldwide web. Cherry County, Nebraska. Late June.

Fungi on the Tallgrass Prairie

Some species of decomposers or or reducers native to the tallgrass prairie were introduced to represent one of the four groups of organisms from the perspectve of the range ecosystem. Decomposers (= Reducers)- In the classic concept of the range ecosystem all biota are classified or grouped by function or role in energy or nutrient transfer as either: 1) producers (green plants fixing energy by photosynthesis), 2) consumers which consume producers either directly (primary consumers which  are herbivores) or indirectly by feeding on herbivores (secondary, tertiary, etc. consumers which are carnivores). Omnivores are those consumers which  function at various times as herbivores and/or carnivores (eg. man) 3) decomposers (= reducers) which are organisms feeding on dead material of producers and/or consumers), and 4) manipulators (= managers)- Man as range manager, forester, wildlife manager, farmer, fisherman, miner. Humans are obviously consumers, but they are unique in their capacity to radically alter ecosystems in ways and at scales that no other organism can (eg. nuclear energy). Producers are autotrophs (autotrophic: self-nourishing) while consumers (and manipulators) and decomposers are heterotrophs (heterotrophic: not self-nourishing but feeding on bodies of producers or consumers and hence dependent on producers as are consumers). Decomposer comprise the least obvious or most arbitrary biotic group. The distinction between decomposing (“feeding on”) dead protoplasm (tissue or organic material) versus consuming live protoplasm becomes unclear with carrion feeders which can range from top order carnivorous predators opportunistically feeding on dead primary consumers (eg. canines, bears), burying and dung beetles, bacteria, and saprophytic vascular plants and fungi. Odum (1971, p.11) found it “… preferable not to designate any particular organisms as ‘decomposers’ but to consider ‘decomposition’ as a process involving all of the biota and abiotic processes as well”. Either way the range student should understand that saprophytes (plants living off of dead or decaying organic matter) are a group essential to functioning of ecosystems through both the one way transfer of energy and the cycling of minerals (= nutrients) within ecosystems. One of the major groups of saprophytes are the fungi (hairs need not be split over the taxonomic treatment of fungi as a kingdom distinct from Plantea, vascular plants, in the five kingdom scheme). Two species of saprophytic fungi were included here. A species of saprophytic vascular plant, a dicotyledon, found in the Sierra Nevada Mixed Conifer Type was included with slides of that forest range cover type (Forest and Woodland Biome slide).

324. Dead man’s hand (Scleroderma geaster)- This earthball or false earthstar or false puffball consist of a tough skin which when it splits apart appears as characteristic thick rays with a conspicuous spore mass in the center. Like the puffballs, this fungus is a member of the Gasteromycetes (stomach fungi) class in the Basidoiomycetes.This large species (knife is 3 1/4 inches) occurs on tallgrass prairies and oak-hickory savannas. It appears in late winter to early spring. Newton County, Missouri, April.

325. Meadow puffball (Lycoperdon cyathiforme or L. caelatum= Calvatia cyathiformis or C. caelata)- Thiis fleshy saprophytic fungus is a member of the Basidiomycetes group. It is common in late summer and early autumn on both native grasslands and permanent introduced pastures where when mature it produces a seemingless stream of spores. Ottawa County, Oklahoma, September.

Fire on Tallgrass Prairie (A Flash Fire of a lesson)

326. All real rangemen love to burn- Fire was a natural component of grassland ecosystems long before man appeared on Earth. Fires which were ignited by natural sources, especially lightening, occurred for millinia before evolution of thinking man (Homo sapiens). Fires ignited by lightening are as much an atmospheric phenomenon and component of climate that determines structure, function, and productivity of grasslands as are precipitation patterns (including drought), temperature regimes, wind, etc. Prehistoric men took their lessons from Nature and fired their ranges in order to produce more feed for game, facilitate hunting and travel, encourage growth of medicinal and pot herbs, and even as a tool of war against enemy tribes. In the Americas white men adopted this practice of the Indians but were careless and excessive in the practice. Thus the first professionally trained range and forest practitioners over-reacted against the otherwise wise use of this essential environmental (largely climatic) component as a tool in Range Management and Forestry. Understandingly this lead to the unwise policy of fire suppression in the name of conservation. Campaigns initiated to stem overburning “backfired” and grasslands were invaded by trees and shrubs, especially when combined with overgrazing. We are slowly relearning the lessons taught by our aboriginal brothers and using prescribed burning as a management practice. Ranchmen in the marvelous “cow country” of the Flint Hills-Osage and Cherokee Prairies Region of Kansas and Oklahoma never forgot the teachings of the Indians. Spring burns are a routine practice among progressive prairiemen. A prescribed spring burn on an excellent big bulestem-upland switchgrass range in the Osage Questas of the Central Lowland Prairie Province. Greenwood County, Kansas. April. Excellent general sources for role of fire on vegetation and in Range Management and Forestry include such standards as Kozlowski and Ahlgren (1974), Wright and Bailey (1982), Chandler et al. (1983), Biswell (1989), Whelan (1995), Bond and van Wilgen (1996), and DeBano et al. (1998). The definitive work on fire and grasslands is Daubenmire (1968). Fire in the central North American grasslands was discussed in the symposium procedings edited by Collins and Wallace (1990). The definitive summary of fire as an ecological factor in range ecology remains chapter 9 in Humphrey (1962; ps. 148-189) while the classic for use of fire by man in agricultural practice is Sauer, (1952; ps. 10-18) and for general grassland management, Sauer (1950). Man’s use of fire in an historic context was covered comprehensively in the series by Pyne (1982, 1984, 1991)  "I know of no basis for a climatic grassland climax but only of a fire grass ‘climax’ for soils permitting deep rooting." (Sauer, 1950, p.  ).

327. Growth of bottomland switchgrass 14 days post burn- Erath County, Texas, March.

328. Growth of bottomland switchgrass 31 days post burn- Erath County, Texas, April.

329. "The only thing that makes me happier than fire is water"- Dense stand of big bluestem on a Flint Hills range about five months following a spring (April) prescribed burn. On the left (to left of conspicuous Baldwin ironweed in center) was an experimental plot that had been treated by repeated spring burns every third year for a number of yeards prior to the current growing season prescribed fire. Many of the tillers of big bluestem had developed into sexually reproductive (flowering) shoots in response to this current growing season burn that was part of the every-third-year burning fire regime. Current and regular periodic spring fires served to elicit this morphological-physiological response. Increased incidence of flowering along with greater density of shoots and height of individual shoots following spring fire is a typical response of big bluestem on tallgrass prairie (Knapp et al. 1998, ps. 203-205). On the right (right of ironweed) another plot had been fired in the same April burn (five months prior to photograph), but this portion of the big bluestem-dominated tallgrass prairie had not been subjected to fire for the last consecutive 15 years (growing seasons). By contrast, big bluestem on this experimental plot did not respond (at least not as greatly) to the spring prescribed fire (ie. there was no increase in flowering, shoot density, or shoot height). This is also a typical result to grassland fire for this climax vegetation. It is apparently the entire fire regime (pattern, including chronology, of burning as well as season, intensity, kind of fire, etc.) and not just conditions of the current fire that is responsible for prairie plant responses to burning. Konza Prairie, Riley County, Kansas. Early August.

330. "Fire? It's about time!" (Or "You're years late")- Stand of big bluestem on the same Flint Hills range and burned by the same April fire as some of the prairie vegetation presented in the immediately preceding photograph (that to left of ironweed) except that this experimental area had not been burnt for 15 consecutived years prior to the current growing season prescribed fire (five months previous). Big bluestem did not respond to the same degree with regard to shoot flowering, density, and height as did big bluestem in adjoining plots that had received spring (April) burns every three years. However, shoot response, especially flowering, was greater than that under annual burning (the every year for eight consecutive years treatment). Again, it is the complete history or fire regime that is important in determinig response of grassland plants to any given fire. Konza Prairie, Riley County, Kansas. Early August.

331. "Another fire? So what else is new?"- Upland big bluestem-dominated tallgrass prairie range that had received prescription fires for the last eight consecutive springs. This was another experimental plot adjacent to the plots ahown and discussed in the two immediately preceding slides. On this range plot there was no increase in sexual reproduction by big bluestem due to fire (there was very little flowering at all). Furthermore, biomass production was about the same between this annually burned tallgrass prairie and that of companion plots that were burnt only one year every 15 years. The main difference between such extreme fire treatments was in flowering rate (proportion of sexually reproductive shoots) of big bluestem and general appearance of the vegetation (less standing and down detritus from previous years on the plot having yearly prescribed fire). Synopsis of burning "shedule": 1) burn every year (as for eight years straight) there is no increase in shoot flowering, density, or height; 2) burn irregularily (as only one year in 15) results in some increase of flowering (development of asexual shoots into sexual shoots), but not as much as at closer interval (say, every third spring) while herbage yield is similar to that under annual burning; 3) burn at regular multi-year spaced intervals (at, in this trial, every third year) there was a big increase in shoot density, height, and flowering rate. Konza Prairie, Riley County, Kansas. Early August.

332. Blackjack oaks top-killed by a hot heading fire on a big bluestem-dominated loamy prairie range site. Blackjacks are resprouting (weakly) from the up7er roots and stumps but continued prescribed burning at intervals of three to five years will reduce oak cover and maintain the fire-type tallgrass prairie in the Cross Timbers Region. Five weeks post burn. Tallgrass Prairie Preserve of The Nature Conservancy (but it was the stewardship of the previous owner, the famed Chapman-Barnard Ranch, whose management  preserved the excellent range condition of this pristine grassland). Osage County, Oklahoma, May.

Natural Revegetation of Tallgrass Prairie

Shown in the following short section were two examples of ongoing natural restoration of tallgrass prairie, recovery of the native prairie plant community through secondary plant succession. One of these recovering prairie ranges was on a bottomland range site while the other was on an upland range site. Grassland vegetation on bottomland had progressed along its sere to climax whereas the upland prairie plant community was roughly in mid-sere with climax grass species just becoming established. Both of these short photographic case histories were on old fields or "go-back land" that had been farmed (row crops and small grains on both fields) for several decades (both for at least a half century) and then retired from farming. The old fields were then abandoned for a period of about 40 years on bottomland and 20-25 years on the upland prairie site. (Obviously there were two different potential climaxes or terminal range plant communities.) No attempts at artificial revegetation such as reseeding had been made on either of the old fields. The bottomland had been mowed for hay once annually since shortly after abandoment. (No comment necessary to describe forage quality--or lack thereof--in early years of vegetation recovery.) The old field on the upland site was mowed annually for hay only a few times afterwhich it was completed "idled" with grazing limited to native animals. The author grew up with and watched land use and recovery of range vegetation on both of these old fields over the course of his lifetime. No systematic records were kept because the author was not present at all times and therefore could not keep accurate or complete notes on management. The following photographs with empherical and incomplete observations were examples of what F.E. Clements called "dynamic vegetation", plant community development by the set of processes known as plant succession (secondary succession in these two instances). These lessons in old field succession were Vegetation Science applied to Range Management as taught in Nature's own lecture hall. Some fellow rangemen and students of vegetation have not so been so fortunte as to have seen such successional sagas unfold. Ya'll will have to be satisfied with this personal and partial narrative, at least until your're better paid.

333. Its reign regained- An old field that through secondary plant succession returned to climax vegetation of mesic tallgrass prairie in the western Ozark (Springfield) Plateau roughly 40 years following cessation of farming. The dominant--far and away--on this Bottomland range site was eastern gamagrass with secondary, though abundant, species including blackeyed susan (Rudbeckia hirta), rosinweed (Silphium integrifolium), purpletop, common wild four o'clock (Mirabilis nyctaginea), and a few seedlings of black walnut (Juglans nigra). When in high school the author helped farm this ground seated on a John Deere Model A. He hauled pea and millet hay (a catch crop planted in a summer drought) off of this little field 45 years before he took these photographs. This land was farmed for a period of about five to seven years after that at which time it was retired from farming by the old couple who owned it. . The field had been mowed once each summer after farming ceased with the first yers hay crop consisting of crabgrass and annual weeds (cows had nothing better to do than pick out grass according to the elderly owners) Annual mowing in late summer has continued ever since tillage ceased. There were no management inputs except for hay harvest. This author first saw eastern gamagrass on the old field approximately 30 years following abandonment. The recovered grassland vegetation shown here was as it appeared about 40 years since the last pass with the plow. A most remarkable story of natural revegetation through secondary succession. Where did the caryopses of this granddad, decreaser grass come from? This little field was on the outer (upper) floodplain of a typical Ozark creek. A climax sugar maple (Acer saccharum)-basswood (Tilia americana)-white ash (Fraxinus americana)-pignut hickory (Carya cordiformis) forest that developed on the north slope of bluffs along the creek was only 70-80 yards from the range plant community shown here. There was no eastern gamagrass in that north slope mesic forest. Likewise, the author was unaware aof any plants of eastern gamagrass upstream from this field. A few other plants of eastern gamagrass were growing in the fencerow across a county "dirt road" from the recovered old field-vegetation presented above. What was their source" Was it the same as that of the ones on this abandoned field or did those across the road come from grain disseminated from this old field stand? So where did the initial Tripsacum dactyloides germinules of recovery emanate from? What where these plant propagules?. The first question above implied that grain was the source of this regenerated stand of eastern gamagrass. The more recently established (the younger) individual clumps of eastern gamagrass appeared to have originated from the initial plants established earlier in ssuccession, and probably by grain disseminated short distances. But what (and where from) were the first disseminules that led to establishment of eastern gamagrass in the revegetation process? Caryopses? Perhaps grains in the soil seed bank that had laid dormant decades. Or could they have been rhizomes, those big, scaly rootstocks of this grandpaw grass species? If initial reestablishment was by asexual reproduction (as from rhizomes) whinther these propagules? Obviously God alone knows. It was a miraculous recovery of a bottomland tallgrass prairie. That much is known by those with their imperfect knowledge. Or perhaps not miraculous at all (except in the sense that a sunrise or birth of a new calf are miracles), but rather the inevitable return of a "super organism" inexorably developing along its sere with the particulars of plant migration yet to be discovered by those who follow F.E. Clements. Trees in the background were those of a bottomland forest adjoining a typical Ozark creek. Many of them were sycamore (Platanus occidentalis), honey locust (Gleditsia triacanthos), white ash, and pignut or bittrnut hickory immediately upstream from the sugar maple-basswood-white ash community that developed on bluffs above the creek. There was certainly no plants of eastern gamagrass or rosinweed in these forest communities. The thin, buff-colored strip of vegetation between creek bottom forest and eastern gamagrass-dominated grassland was naturalized tall fescue (Festuca arundinacea). This agronomic species has naturalized over much of the Ozark Region and is frequently a noxious species especially on tallgrass hay meadows and pastures. Tall fescue stood no chance against eastern gamagrass and, in fact, the latter was slowly encroaching on the stand of tall fescue. Sic 'em grandpaw! Ottawa County, Oklahoma; greater floodplain of Modoc Creek. July (estival aspect); peak bloom in the most common composite species and grain-ripe to grain-shatter in eastern gamagrass.

334. Throne reclaimed- Stand of eastern gamagrass with rosinweed as the local assocaite species on an old farm field that revegetated through secondary plant succession to a climax, bottomland tallgrass prairie in the Ozark Plateau. This was another view of the same old field and natural restoration of potential natural range vegetation introduced in the preceding two photographs and described thereunder. The creek bottom forest and narrow strip of tall fescue behind the eastern gamagrass and rosinweed was also described immediately above. It bore repeating that eastern gamagrass was slowly invading the established sward of the naaturalized, often weedy, tall fescue. (That vegetational development also makes frequent visits to this recovered tallgrass prairie range well worth the short excursions.) Ottawa County, Oklahoma; greater floodplain of Modoc Creek. July (estival aspect); peak bloom in rosinweed and grain-ripe to grain-shatter in eastern gamagrass.

335. Regaining its ground- The climax regime of tallgrasses (here represented by big bluestem and upland switchgrass) was slowly--perhaps inexorably--returning to an old field 20-25 years post abandonment following at least 60 years of farming. In fact, this land could have been farmed for as long as 70 or even 80 years, but there was living locals who could remember back to that time. This old field had produced various crops including wheat (Triticum aestivum), dent corn ( Zea mays var. indentata), grain sorghum (Sorghum vulgare= S. bicolor), and soybeans (Glycine max) for at least the better part of six decades. Even the exact time (year) of the last field crops was apparently forgotten by current owners, but the author who was raised in this locality can trace the point of retirement from cropping from roughly 20 to something less than 30 years prior to time he took these photographs. No attempts at artificial revegetation were made. Seral vegetation on this "go-back land" was mowed for hay one time in each of a few summers (exact number of hay harvests was unknown to the author), but this was soon discontinued. No mowing had taken place on this old field for at five or six years prior to the year in which these photographs were taken This land had passed through the textbook sequence of old field succession from an initial pioneer sere of crabgrass, horseweed, giant ragweed, old field (annual) threeawn, redroot pigweed, and some cocklebur through to the seral community dominated by biennial forbs and weedy or ruderal perennial grasses, especially the native broomsedge bluestem and the naturalized Johnsongrass. The seral plant community shown here and in the succeeding photograph was the latter phase of the weedy, perennial (with lots of anuals and biennials still present) stage that was progresssing into the next (another mixed) stage of weedy perennials grasses plus weedy forbs and with initial establishment (Clementsian migration) of native climax (decreaser) tallgrass species. Big bluestem and switchgrass of the upland form as the first of the Four Horsemen tallgrasses had achieved a successional "beach head" on this "go-back land". Thickening of these individual clumped plants by increasing number of shoots (asexual reproduction) was part of the process Clements called aggregation (Weaver and Clements, 1038, ps. 4, 145-147). This was the second example presented in this section showing natural recovery of tallgrass prairie in the western Ozark Plateau by secondary plant succession. The same question that was asked for the first example presented and described above was reasked here, "Where did the plant propagules of these climax grass species come from?" How did the initial disseminules get here or, if present from the initial virgin sod that was plowed 60 to 80 years previously, how did they persist under decades of farming? What were the germinules? Tiny, fluffy spikelets? Fragments of rhizomes or rootcrowns?. What were the agents of dispersal (wind, birds, rodents, even farm implements perhaps)? Numerous plants of big bluestem were growing within approximately one-fifth mile from these plants, but the author was unaware of any switchgrass within a five mile radius. Imponderables. Broomsedge bluestem was the dominant plant in this mid-sere vegetation. The associate species varied locally and seasonally. For example, in this late spring society the naturalized Eurasian biennial umbel, wild carrot or Queen Anne's lace (Daucus carota), was the major forb and a local associate. Other major species included Johnsongrass (Sorghum halepense), a naturalized perennial grass that was also a local associate, hairy crabgrass (Digitaria sanguinalis), Japanese chess (Bromus japonicus), yellow hop clover (Trifolium agrarium), and tall fescue. Clearly, exotic invaders dominated range vegetation on this old field; it was also obvious that the native decreaser tallgrasses had invaded so that aggregation was underway and migration ongoing. Ottawa County, Oklahoma. Mid-June (vernal aspect); various phenological stages depending on plant species.

336. Winter yet a new beginning on an old field- Scene of the recovering upland tallgrass prairie just shown in the winter following the above summer scenes. Most of the herbage was broomsedge bluestem, but three plants of switchgrass (upland form) were present (foreground to near background). There were even more plants of big bluestem, but the readily degradable shoots of this decreaser had already decomposed to the point of falling to the land surface. Ottawa County, Oklahoma. January (hibernal aspect); depth of winter dormancy.

Nebraska Sandhills (and related postclimax tallgrass types)

Sand and sandy soils (also gravel for that matter) have a higher proportion of macropores (noncapillary pores) so that there is less water runoff and more water that infiltrates this rooting material for plants. Conversely,there is less water retained in the coarse or heavy textured soils as compared to fine (light)-textured soils such as clay; however, , more of the retained water in coarse textures (eg. sand) is available for plant absorption. Also many plants that are adapted to coarse-textured soils have a high proportion of their roots near the land surface so that they absorb this temporarily shallow water before it evaporates or percolates below their root zones. Net result is that the native vegetation on sand, sand dunes, and high sand-content soils is typically more mesophytic than vegetation on adjacent or neighboring soils comprised of less coarse particles (those that have finner soil texture). This was explained in Weaver and Clements (1938, p. 203-204) by the Clementsian concept of chresard. Chresard is that part of total soil water, the whole content of water in soil (= holard), that is available for plant absorption or use by the plant part. The echard (to withold) is that portion of water still present (held or retained) in soil at the permanent wilting point (ie. water that is unavailable for plant use; water that adheres to tightly to soil particles that plants cannot absorb it). Holard minus chresard equals echard (or any other arrangement of these portions ssuch as holard minus echard equals chresarde or chresard plus echard equals holard). In the Clementsian monoclimax model natural vegetation of sandy environments if postclimax "In Nebraska the relation [transition from true to mixed prairie] is further disturbed by the extensive sand-hill region, in which the high chresard favors a postclimax of tall grasses far beyond their proper climate [ie. beyond the climatic or regional climax]" (Weaver and Clements, 1938, p. 524). "In the vast sand-hill area of Nebraska, the tall-grass postclimax attains its best development, which is assumed to reflect the climate when the prairies were occupied by the bluestems and their associates some millions of years ago" (Weaver and Clements, 1938, p. 86). This also included shrubs such as New Jersey tea, sand sagebrush (Artemisia filifolia), and soapweed yucca. The sandhill region of Nebraska and similar smaller areas in other parts of the Great Plains including some in Colorado, Kansas, Oklahoma, Texas, and New Mexico support more mesic (more moisture-requiring) range vegetation than that of less porous soils in the same region or area. In the vernacular of Tansley's polyclimax model or Whittaker's climax pattern interpretation postclimax vegetation would be edaphic and/or topographic climax vegetation (edapho-topographic climaxes). Same ecological (successional or climax) outcome. Tallgrass prairie range vegetation in the Nebraska Sandhills (and some similar range communities in Colorado, Oklahoma, New Mexico, Texas, etc.) is in the Mixed Prairie Region though, of course, it is not mixed prairie. As such, postclimax or edaphic-topographic climax tallgrass prairie was covered in this chapter. This included such range types as sand sagebrush- and shinnery oak (Quercus havardii)-tallgrass savannas as well as the classic Nebraska Sandhills tallgrass prairie. Range types of mixed prairie, including some in the Nebraska Sandhills (not all Sandhills grassland is tallgrass prairie), were (logically) dealt with in the chapter, Mixed Prairie. Organization note: The Nebraska Sandhills as well as related tallgrass prairie subtypes or forms of the Southern Great Plains (= High Plains, Staked Plains, Llano Estacado) were treated separately in the subsequent chapter, Tallgrass Prairie (Interior)- II. A few photographs of the famed Nebraska Sandhills were retained in this chapter to make for a smoother transition to that next chapter that was devoted to tallgrass prairie vegetation of the semiarid zone.

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

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

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

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

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