Tallgrass Prairie

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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).

* Note: In the North American biotic community classification of Brown et al. (1998) all cover types within the tallgrass prairie were encompassed by the generic--and overly broad-- designation of Bluestem "Tall-Grass" Series 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.

3. 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]).

4. 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 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.

5. 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.

6. 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 (eg. 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.

7. 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.

8. 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).

9. Upland tallgrass (bluestem) prairie- Big bluestem was the dominant and prairie dropseed the associate species on this Osage Questa 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 SRM 710 (Bluestem Prairie), probably more specifically.

10. 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).

11. 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.

12. 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.

13. 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).

14. 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).

15. 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).

16. 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).

17. 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).

18. 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).

19. 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).

20. 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).

21. 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 (Missosuri Prairie).

22. 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).

23. 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.

24. 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.

25. 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.

26. 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).

27. 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.

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

29 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!).

30. 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).

31. 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.

32. 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).

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

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

35. 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).

36. 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).

37. 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).

38. 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).

39. 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).

40. 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).

41. 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).

42. 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.

43. 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.

44. 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.

45. 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.

46. 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.

47. 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.

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

49. 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).

50. 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.

51. 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.

52. 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.

53. 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.

54. 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.

55. 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.

56. 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.

57 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).

58. 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.

59. 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.

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

61. 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.

62. 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.

63. 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.

64. 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.

65. 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.

66. 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.

67. 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.)

68. 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.

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

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

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

72. 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.

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

74. 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.

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

76.  Silver bluestem (Andropogon saccharoides= Bothriochloa sacharoides) 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. Erath County, Texas. July.

77. 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.

78. 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.

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

80. 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.

81. 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.

82. 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.

83. 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.

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

85. 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.

86. 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 everywhere else). Grass burrs are a textbook example of zoochory, dispersal of plant disseminules by animals, including rangemen.

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

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

89. 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.

90. 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.

91. 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.

92. 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.

93. 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".

94. 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.

95. 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.

96. 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

97. 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.

98. 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.

99. 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. 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.

100. 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.

101. 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".

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

103. 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.

104. 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.

105. 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.

106. 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.

107. 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.

108. 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.

109. 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.

110. 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.

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

112. 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.

113. 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.

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).

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

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

116. 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.

117. Inflorescences of Maximillian sunflower- Wichita Mountains Refuge.

118. 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.

119. Inflorescence of pale purple coneflower (Ecninacea pallida)- 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.

120. Shoot of white sweetclover- Leaves and inflorescences of white sweetclover. Ottawa County, Oklahoma. June.

121. "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 favorites. 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. Three Solidago species commonly found on tallgrass prairie were included below just to give students a taste of plant speciation of the range.

122. 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.

123. 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.

124. 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.

125. 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.

126. 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.

127. 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.

128. 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.

129. 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.

130. 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.

131. 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.

132. Flowering shoot of butterfly milkweed- Leaves and inflorescence of the milkweed known simply as butterfly weed. Ottawa County, Oklahoma. June.

133. 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.

134. 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.

135. Inflorescence of white snakeroot.

136. 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.

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

138. 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.

139. 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.

140. 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.

141. 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.

142. 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.

143. 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.

144. Indian paintbrush (Castilleja coccinea)- This 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.

145. 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.

146. 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.

147. 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.

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

149. 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.

150. 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”. 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).

151. 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.

152. 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.

153. 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.  ).

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

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

156. Blackjack oaks top-killed by a hot heading fire on a big bluestem-dominated loamy prairie range site. Blackjacks are resprouting (weakly) from the upper 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.

157. 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 601 (Bluestem Prairie).

158. 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 601 (Bluestem Prairie).

159. 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 601 (Bluestem Prairie).

160. Prairie sandreed (Calamovilfa longifolia)- This attractive tallgrass is the dominant species on deep sand sites in the greater Sandhills Region of Nebraska and adjoining areas of Colorado, Kansas, and South Dakota. This is locally a major hay grass and, unlike associated panicoid grasses such as the bluestems, prairie sandreed "cures on the vine" (ie. retains higher nutritive value in dead herbage) much like other eragrostoid species such as the less productive grama grasses and buffalograss. Prairie sandreed is often the key species on many range sites (and ranches). Hays County, Nebraska. July.

161. Postclimax tallgrass prairie- Mixed prairie on deep sand in the Southern High Plains consisting of two layers: 1) a tallgrass overstorey dominated by sand bluestem (Andropogon hallii) with little bluestem and upland switchgrass as associate species and 2) a mid-grass understorey with needle-and-thread and sideoats grama as dominants along with traces of sand dropseed and blue grama. Blue grama is usually thought of (and was treated elsewhere herein as) a shortgrass species, but on this Deep Sand range site it attained a height approaching that of sideoats grama and it was so scarce that it did not comprise a shortgrass layer in this grassland vegetation. Sand sagebrush (Artemesia filifolia) also occurred in trace proportions such that it did not add a shrub layer. This vegetation was thus grassland and not a mixed grass-sagebrush shrub savanna (a form presented immediately below). Forbs were not abundant, but wild alfalfa (Psoralea tenuifolia) was present. Scale of this tallgrass form of mixed prairie that developed in a 17 inch precipitation belt can be guaged by the four-foot barbed wire fence. Shoots of sand bluestem remaining (and broken) from the previous growing season extend above the top wire. The short shoot or "culmless" sand bluestem had just started to elongate it shoots and was still in the pre-boot stage during the current growing season. The rolling sand dune microrelief of this range site was shown conspicuously by featuring a dune in extreme left midground. In the photograph the dune sloped down from left (top of dune) to right (bottom of dune) so that the base level of the dune was directly in front of the second steel post from the left side of the slide. Mixed prairie postclimax grassland dominated by tallgrass species, especially sand bluestem, was discussed in Weaver and Albertson (1956, ps. 261-265, 295-299). The concept of postclimax will be confusing to both beginning students and "climax" students who were not instructed or self-taught in the Clementsian paradigm. It is helpful-- if not essential-- for students to have some grasp of the concept of postclimax (and other aspects of the Clementsian model such as preclimax and disclimax) because so much of the seminal and classical work on range vegetation, especially that of grasslands, was interpreted and described in the language of the Clementsian lexicon. The sand bluestem-dominated mixed prairie range presented here was but one example. Aside from the need to be fluent in the prevailing language in which most of North American vegetation was described, concepts like postclimax remain useful in providing a rational explanation for development and persistence of many North American range types. Postclimax tallgrass prairie and mixed prairie dominated by tallgrass species (like sand bluestem) is one of the textbook examples of this. At regional scale the Sandhills Region of Nebraska and Colorado is one of the largest remaining units of postclimax vegetation while smaller areas of sandhills prairie like that featured in this slide are outliers of postclimax plant communities. Readers were referred to Weaver and Clements (1938, ps. 82-83, 85-86, 102, 110, 462, 475, 520-521). Clements (1936, p. 269) used the sandhills tallgrass as an example of postclimax, but in a pattern characteristic of Clement's writing he did not offer a concise definition of postclimax. Allaby (1998) did: "In the monoclimax model of climax vegetation development, communities differing from the climatic climax, owing to cooler and/or moister conditions than are characteristic of the regional climate". For example, soils of the Sandhills permit faster infiltration rates and greater absorption of precipitation to deeper depths in the soil profiles which allows development of vegetation more mesic than that which is typical for the general region of the High Plains. Grassland vegetation of humid and subhumid regions can develop in a semiarid region due to more mesic soil conditions, a function of coarser-textured soils. Southern High Plains. Cimarron County, Oklahoma. July. FRES No. 38 (Plains Grassland Ecosystem). K-62 (Bluestem-Grama Prairie). SRM 720 (Sand Bluestem-Little Bluestem, Dunes). Deep Sand range site.

162. Sand bluestem-little bluestem-sand sagebrush— A sand dune cover type. Clements and Weaver (1938) regarded such communities as postclimax, a tallgrass variant type in the mixed prairie region. This cover type is shown in the grassland section because it is a grassland type and is not a sand sagebrush-sand bluestem savanna. This type occurs exclusively on deep sand sites and is interspersed among blue grama-buffalograss, western wheatgrass, and other forms of mixed prairie and shortgrass plains grasslands.  Chaves County, New Mexico.  June.  FRES No. 39 (Prairie Ecosystem). K-63 (Sand Sagebrush-Bluestem Prairie). SRM 720 (Sand Bluestem-Little Bluestem, Dunes).

163. Coastal prairies and marsh vegetational area of Texas- a landscape mosaic of many ecosystems from estuaries to tallgrass prairie to patches or corridors of the "running" form of live oak (Quercus virginiana) and red bay (Persea borbonia). San Antonio Bay on Aransas National Wildlife Refuge, winter home of the whooping crane (Crus americana).Coming inland from the bay the zones of various communities (many of them consociations or populations) are: 1. smooth cordgrass (Spartina alternifolia), salt marsh, 2. a marsh commuity of saltmarsh bullrush (Scirpus californicus, S. americanus), black needle rush or slender spike rush (Eleocharis acicularis), and whitetop sedge (Dichromena colorata), salt marsh, 3. gulf and marshhay cordgrass (Spartina spartinae, S. patens), tidal flat, and 4. live oak – red bay shrub community in the foreground. Refugio County, Texas. Late heimal- prevernal aspect, February. SRM 726 (Cordgrass) of several forms and variants. Beardgrass Series (Gulf Coastal Grassland) of Brown et al. (1998). Western Gulf Coastal Plains- Mid-Coast Barrier Islands and Coastal Marshes Ecoregion, 34h (Griffith et al., 2004).

164. Texas Coastal Praries and Marshes- Zones of vegetation from San Antonio Bay. Coming from the bay inland these are: the short form of smooth cordgrass (tan-colored zone on far side of the bay) ,  the green zone is saltmarsh bulrush (Scirpus maritimus) and black needlerush (Juncus      roemerianus),  the zone of low-stature vegetation inland from the green zone is the grass community dominated by seashore saltgrass, shoregrass, sea blite, and Virginia glasswort or saltwort,  the zone of tufted plants is a tidal flat dominated by the cespitose marshhay cordgrass with sea ox-eye plus some scattered Gulf cordgrass, and  the inland-most or woody vegetation (foreground) is live oak-red bay (Persea borbonia)-yaupon forest.  The two zones of 1) smooth coregrass and 2) saltmarsh bulrush-black needlerush comprise saltmarsh whereas the 3) salt-shoregrass community and the 4) marshhay cordgrass make up the tidal flat.  Vegetation of these herbaceous zones were shown in preceding slides. Calhoun County, Texas.  Hibernal aspect, February. No single FRES or K-unit could describe this. Texas Coastal Prairies and Marshes Vegetation (= Land Resource) Area. Beardgrass Series of Brown et al. (1998). Western Gulf Coastal Plains- Mid-Coast Barrier Islands and Coastal Marshes Ecoregion, 34h (Griffith et al., 2004).

165. Texas Coastal Bend Prairie- Arch-typical Texas coastal prairie that is basically a consocition of seacoast little bluestem (Andropogon littoralis= Schizachyrium scoparium var. littoralis) with silver bluestem as the major associate and smaller proportions of such tallgrass species as crinkleawn (Trachypogon secundus), Paspalum species, Gulf cordgrass, Indiangrass, and Pan American balsamscale (Elyonurus tripsacoides). Almost no forbs; no woody plants. Aransas National Wildlife Reguge, Refugio County, Texas. October. FRES No.39 (Prairie Ecosystem), K-69 (Bluestem-Sacahuiste [Gulf Cordgrass] Prairie), SRM 711 (Bluestem-Gulf Cordgrass Prairie). Beardgrass Series of Brown et al. (1998). Western Gulf Coastal Plains- Mid-Coast Barrier Islands and Coastal Marshes Ecoregion, 34h (Griffith et al., 2004).

166. Upland (versus bottomland or beach) coastal prairie- This scene is a transect shot extending from a few feet above sea level down literally into the Gulf waters of San Antonio Bay. Several range sites occur along the landscape seen here which extends from coastal prairie to the salt water marsh of smooth cordgrass (spartina alterniflora) out in the bay. Species visible in the prairie of the foreground include seacoast bluestem (the most common species), big bluestem, silver bluestem, Indiangrass, bottomland switchgrass (ie. a Four Horsemen prairie the same as the Flint Hills of Kansas, Osage Hills of Oklahoma, Sand Hills of Nebraska, or Blackland Prairie of Texas), Pan American balsamscale, common reed (Phragimites communis), bushy bluestem (Andropogon glomeratus) withTurk's cap and Gulf Coast coneflower as major forbs.

167. Pan American balsamscale- This member of the Andropogoneae tribe provides forage of only fair to, rarely, good nutritive value but it is a common associate on coastal prairies. It is a distinctive grass but a species lesser known to most rangemen. It was included here to illustrate the vast diversity of species on North American grasslands.

168. Inflorescence of Pan American balsamscale.

169. Composite shot of both the coastal prairie and Gulf salt marsh- The foreground is a tallgrass prairie with bottomland switchgrass conspicuous at far right along with bushy bluestem, seacoast bluestem, the conspicuous Gulf Coast Mexican hat, and common reed (visible as the tallest three or four rank shoots at front edge of water). The background beginning at back edge of the stream is a smooth cordgrass salt marsh. Two distinct range cover types (and two distinct Kuchler units and FRES Ecosystems) are clearly separated by the water line: brackish or salt water from San Antonio Bay reaches up to the back side of the stream (which is mostly fresh water) and supports smooth cordgrass which is Gulf Coast salt marsh: FRES No. 41 (Wet Grasslands), K- 70 (Southern Cordgrass Prairie), SRM  806 (Gulf Coast Salt Marsh). In front of the fresh water stream which serves as a barrier against intrusion of Gulf salt water from the bay is coastal tallgrass prairie: FRES No. 39 (Prairie Ecosystem), either K- 66 (Bluestem Prairie) or K-69 (Bluestem-Sacahuiste [Gulf Cordgrass] prairie and thus either SRM 710 (Bluestem Prairie) or 711 (Bluestem-Gulf Cordgrass Prairie). Beardgrass Series of Brown et al. (1998). Western Gulf Coastal Plains- Mid-Coast Barrier Islands and Coastal Marshes Ecosystem, 34h (Griffith et al., 2004). There is very little of either marshhay or Gulf cordgrass so it is basically tallgrass prairie as explained in the previous landscape scene. Aransas National Wildlife Refuge, Refugio County, Texas. October.

170. Gulf Coast saltwater cordgrass marsh- This unique grassland community is a consociation of smooth cordgrass (Spartina alterniflora) that extends the entire length of the Gulf Coast clear up along the Atlantic Seaboard, often in patches or of intermittent distribution, to the salt marshes of New England extending to Nova Scotia. These regularly flooded tidal or Gulf marshes  “… are almost exclusively Spartina alterniflora marshes…” and are “… almost a natural monoculture of  Spartina alterniflora …” (Teal, 1986, p.1). It forms the Great Barnstable Marsh at Cape Cod, Massachusetts and the smooth cordgrass marshes at Cape Hatteras National Seashore, South Carolina. Furthermore the S. alterniflora marshes have a worldwide distribution occurring in Argentina and western Europe, including the British Isles (Teal, 1986, p.3). It has been widely studied and shown that there are two varieties of S. alterniflora: tall form and a short form of smooth cordgrass with the tall being on tidal creeks with fresh water and the short variety on sites or local areas subject to flooding with salt water. Some workers reported an intermediate form between fresh (or slightly brackish) and salt water (Haines and Dunn in Chabot and Mooney, 1985, ps. 323-341).   This plant community is, of course, a major part of the estuaries all along the coast. In fact, smooth cordgrass is often the major producer in these estuaries (Odum, 1971, p. 357). It was while working with these tidal marshes in Georgia that Eugene Odum discovered that the salt marsh-estuary ecosystem (the tidal or coastal wetlands= coastal tidelands) is probably the most productive ecosystem on Earth. Work like this and that by Teal (and the very popular book by Teal and Teal [1969]) as well as publicity by prominent conservationists like Rachel Carson spawned a nationwide effort to save what remained of United States’ ocean marshes and estuaries. The story of this largely successful, remarkable conservation program was published by (Siry, 1984).  Of more direct historical interest to range students is the fact that the smooth cordgrass salt marshes formed part of the vast coastal and bay range that was the cradle of the range cattle industry in Texas and Louisiana. The famous Longhorns in this region were appropriately dubbed by Texas cowmen as "coasters" and "sea lions". FRES No. 41 (Wet Grasslands Ecosystem), but Kuchler did not map the smooth cordgrass tidal flats (tidal, coastal, or sea marshes). Specifically the smooth cordgrass "monoculture" is one component or varient of SRM 806 (Gulf Coast Salt Marsh). Beardgrass Series of Brown et al. (1998). Western Gulf Coastal Plains- Mid-Coast Barrier Islands and Coastal Marshes Ecoregion, 34h (Griffith et al., 2004).

171. Inside a cordgrass (gulf and marshhay) tidal flat— A woody component of the composite Baccharis halimifolia and the borage, seaside heliotrope (Heliotropium curassavicum ), surrounded by red bay – live oak shrub (or actually trees) to constitute a “forest” of sorts. Aransas Refuge, Refugio County, Texas. FRES No. 41 (Wet Grasslands Ecosystem). K-70 (Southern Cordgrass Prairie). SRM 726 (Cordgrass) or, partially , SRM 719 (Mesquite-Live Oak- Seacoast Bluestem). Beardgrass Series of Brown et al. (1998). Western Gulf Coastal Plains- Mid-Coast Barrier Islands and Coastal Marshes Ecoregion, 34h (Griffith et al., 2004).

172. Aransas Tidal flat range community- Marshhay cordgrass (green tufts on right margin), seashore saltgrass (adjacent and immediately left of cordgrass), shoregrass (Monanthochloe littoalis) left and adjacent to saltgrass, sea ox-eye (Borrichia frutescens) is the pea-green forb, common brown forb producing most cover is perennial sea blite (Suaeda conferta), and scattered yellow forbs are Virginia glasswort or saltwort (Salicornia virginica).

173. Species of Aransas tidal flats- Annual saltwort (S. bigelovii) in shoregrass.

174.  Sward of seashore saltgrass and shoregrass.

175. Seashore paspalum (Paspalum vaginatum)- This species sends stolons up to edge of (frequently extending into) Gulf water. It is one of the most salt water-tolerant grasses in the world.

176. Archtypical scene of coastal prairie— Swale-and-ridge gulf grassland comprised of the Four Horsemen tallgrasses and surrounded by live oak mottes plus laurel oak (Quercus hemisphaerica), red bay, and yaupon (Ilex vomitoria). Associate grassland species include marshhay cordgrass and bushy bluestem (Andropogon glomeratus). Much of the little buestem is the sea coast form treated by some as a variety of little bluestem (Schizachyrium scoparium var. littoralis) but as a distinct species (Andropogon littoralis)by Hitchcock and Chase (1950).Aransas National Refuge, Refugio County Texas. Prevernal aspect, February. FRES No. 41 (Wet Grasslands Ecosystem) and FRES No. 16 (Oak-Gum-Cypress Ecosystem). K-70 (Southern Cordgrass Prairie) surrounded by K-81 (Live Oak-Sea Oats). SRM 726 (Cordgrass) and SRM 719 (Mesquite-Live Oak-Seacoast Bluestem). Beardgrass Series of Brown et al. (1998). Western Gulf Coastal Plains- Mid-Coast Barrier Islands and Coastal Marshes Ecoregion, 34h (Griffith et al., 2004).

177. Closer view of the fauna of the Scirpus colony- A female American alligator (Alligator mississippiensis) is the representative of the largest reptile of the Atlantic coastal prairies and marshes. The ‘gator was reportedly 13 feet long. FRES No. 41 (Wet Grasslands Ecosystem), K- 42 (Tule Marshes), SRM 807 (Gulf Coast Fresh Marsh).

178. Cordgrass salt flats of the Gulf coastal prairies- This pristine, verdant sward is a strikingly beautiful consociation of Gulf cordgrass (Spartina spartinae). This climax grassland was one of the most extensive range communities in the virgin coastal prairies and marshes of the Atlantic Gulf, especially in Texas and Louisiana. It formed vast essentially single-species stands along the coastal salt flats and inland sites that are subject to flooding or intrusion of salt water. Gulf cordgrass is tolerant of submersion and can extend to beaches. The excellent range seen here is maintained by routine prescribed burning and proper grazing management. It is a picture- perfect example of perfect stewardship.   Ranch, San Patricio County, Texas. October. FRES No. 41 (Wet Grasslands Ecosystem), K-70 (Southern Cordgrass Prairie), SRM 726 or FRES No. 39 (Prairie Ecosystem), K-69 (Bluestem-Sacahuiste [Gulf Cordgrass] Prairie), SRM 711 (Bluestem-Gulf Cordgrass Prairie): see explanation under next slide. Western Gulf Coastal Plains- Southern Subhumid Gulf Coastal Prairies Ecoregion, 34b (Griffith et al., 2004).         "The sight of a native turf, whether of shortgrass carpeting the Earth or tallgrass waving in the wind,  restores my soul."  (J. Frank Dobie).

179. Gulf cordgrass- This cespitose species forms large, sometimes massive, clumps that make travel in vehicles uncomfortable but which provides abundant and good to excellent forage when young. Communities dominated— often populated almost exclusively —by Gulf cordgrass furnished the feed for the earliest days of cattle ranching in Texas and Louisiana. Jordan (1981) proved that cattle ranching in Texas originated from Anglo Saxon roots in South Carolina and began along the Gulf Coast and NOT as traditionally taught in the Rio Grande Plains from horseback Hispanic ancestry. Gulf cordgrass  salt flat prairies— the few that remain —still provide excellent natural pasture for large ruminants as well as being part of the coastal habitat for the endangered Atwater prairie chicken. O’Conner Ranch, San Patricio County, Texas. October. FRES No. 41 (Wet Grasslands), K-70 (Southern Cordgrass Prairie), SRM 726 or FRES No. 39 (Prairie Ecosystem), K-69 (Bluestem-Sacahuiste [Gulf Cordgrass] Prairie), SRM 711 (Bluestem-Gulf Cordgrass Prairie). There are characteristics and components of both of these depending on if land is covered with standing water and/or soil saturated for long periods or if it has features more like those of drier upland prairie.Generally, the Beardgrass Series of Brown et al. (1998). Western Gulf Coastal Plains- Southern Subhumid Gulf Coastal Prairies Ecoregion, 34b (Grifith et al., 2004).

180. Inflorescence of Gulf cordgrass-  O'Connor Ranch, San Patricio County, Texas. October.

181. Sea oats-seacoast bluestem coastal prairie- An interrupted narrow strip of tallgrass prairie, a dry prairie (versus wet prairie and marsh), sporadically lines the Gulf Coast. This unique form of the vast tallgrass-dominated grassland in the heart of North America is the coastal terminus of a belt of humid climate-derived prairie that once extended southward from Manitoba to the Gulf. Some of the same dominant panicoid species extended throughout the latitude of this vast (once vast; now largely plowed or paved over) grassland. One of the most consistent of these species was little bluestem and its closely related taxa (both ecotypes and taxonomic varieties that have often been interpreted as separate species). The Gulf Coast "version" of little bulestem which is the dominant species of the coastal tallgrass prairie is sea coast bluestem (Andropogon littoralis, to most early day American agrostologists like Hitchcock, and Schizachyrium scoparium var. littoralis to recent authorities like Gould). The distinguishing species of this unique form of bluestem prairie (ie. the key ecological species giving this vegetation it's distinctive quality, and it's designation) is sea oats (Uniola paniculata). The Society for Range Management (Shiflet, 1994) designated and described this rangeland cover type as the sea oats prairie. Sea oats were shown here in full-flower on low blow-out sand dunes that were stabilized by the roots and rhizomes of this eragrostoid grass. The spikelets of sea oats are probably the most laterally compressed of any North American grass and their beauty in the massive inflorescences provides as distinctive a quality as the picturesque grassland itself. The view here is westward toward the mainland from this barrier island. Immediately behind this stand of sea oats is a wind-tidal flat formed by the combination of high heat, relatively constant temperature, persistent winds carrying salt particles, and saline soils. Wind-tidal and -deflation flats, deflation troughs, dunes of several kinds (eg. coppice, barchan, transverse, blow-out) and other miscellaneous local landforms create a patchwork of plant communities which are various seral stages in plant succession that probably terminates in a seacoast bluestem-sea oats climax.. Students of Plant Ecology should recall that many of the pioneering studies of plant succession in both America and Europe were done on sand dunes, the most famous of which was that of H.C. Cowles from which F.E.Clements took his grand model of "dynamic vegetation". In the far background is an interdunal area dominated by seacoast bluestem as shown in detail in the succeeding slide. Padre Island National Seashore, Texas. Hibernal aspect, February. FRES No. 39 (Prairie Ecosystem). Mapped by Kuchler as K-70 (Southern Cordgrass Prairie) which indicated that it was part of FRES No. 41 (Wet Grassland Ecosystem). But, as was the case for several North American range types, Kuchler mapping scale was too coarse (large or general) to include the narrow belt of dry prairie dominated by seacoast bluestem and sea oats within the more general coastal marsh and wet prairie zone. More precisely this is an outlier of K-66 or K-69 (a form of Bluestem Prairie). SRM 723 (Sea Oats). Part of the overly broad Beardgrass Series of Brown et al. (1998). Western Gulf Coastal Plains- Laguna Madre Barrier Islands and Coastal Marshes Ecoregion, 34i (Griffith et al., 2004).

182. Seacoast bluestem-sea oats prairie- This community most likely represents the potential natural terminal vegetation of the barrier islands and high dune zone of mainland beaches. Seen here is the stabilized blow-out dune zone which is the interior grassland of the barrier island. Foredunes just beyond the swash zone of the beach and wind-deflation flats and back-island dunes behind the flats (mainland-ward) act as borders and barriers to this seacoast bluestem-sea oats climax grassland. The sea oats prairie, as it has been referred to (eg. Shiflet, 1994) based on the distinctiveness and specificity of this sand-binding rhizomatous grass, is found most consistently on the North American barrier islands such as Padre and Mustang Islands along the Texas coast. Vegetation of these barrier islands is complex and occurs as zones, the community compositions of which are determined by distance from the beach and strand (moving inland) or from the mainland (moving seaward or toward the coast). This example was found on north Padre Island with the extremely salty Laguna Madre and its back-island dunes behind these stabilized blow-out dunes. These dunes were populated primarily by the two dominant species with sea oats dominant on dune crests and seacoast bluestem dominant lower on the dune faces and the ground among adjacent dunes. Within the overall seacoast bluestem-sea oats prairie there are numerous local sites (and even microsites) such as small fresh water or brackish marshes, blowouts, and deflation flats. All of these are various seral stages in the xerosere of the island with the stabilized sand dune and interdunal flats comprising the local climax grassland vegetation of which the dominants are typically sea coast bluestem and sea oats with such associate grasses as gulfdune paspalum (Paspalum monostachyum), bitter panicgrass (Panicum amarum), and, locally, big bluestem. Prominent forbs include whitestem wild indigo (Baptisia laevicaulis), largeleaf pennywort (Hydrocotyle bonariensis), and morningglory species (eg. Ipomoea stolonifera). All examples of vegetation presented in the following slides are of this general climax community. The relatively flat (at least less choppy) land surface seen here in the foreground was largely a stand of seacoast bluestem with whitestem wild indigo the apparent associate species. Bitter panicgrass was also commonly interspersed around clumps of the cespitose bluestem. Padre Island National Seashore, Texas. Hibernal aspect, February. FRES No. 39 (Prairie Ecosystem). Mapped by Kuchler as K-70 (Southern Cordgrass Prairie) which indicated it was part of FRES No. 41 (Wet Grassland Ecosystem), but, as was found for several North American range types, Kuchler mapping scale was too large or too general to include the narrow belt of dry prairie dominated by seacoast bluestem and sea oats within the more general coastal marsh and wet prairie zone. More precisely this is an "island" of K-66 or K-69 (a form of Bluestem Prairie). SRM 723 (Sea Oats). Beardgrass Series of Brown et al. (1998). Western Gulf Coastal Plains-Laguna Madre Barrier Islands and Coastal Marshes Ecoregion, 34i (Griffith et al., 2004).

183. Stabilized blow-out dune in climax grassland- Here in the interior of Padre Island, seacoast bluestem was the local dominant (with sea oats the associate species) atop a blow-out dune. With a slight shift in wind direction or with extended periods of higher wind speeds dune tops develop small blowouts which are covered by sea oats. Sea oats also quickly populate some of the more recent sand dunes and function as both a pioneer and a climax species (at least at local scale). The bare branches of the shrub extending above the dune in the center are of the deciduous black willow (Salix nigra). Sand dunes on the barrier islands frequently have persistently wet or moist layers of sand below the land surface that enables hydrophytes such as willow to persist in an otherwise xeric environment. Also, sand usually has high infiltration rates allowing dunes to absorb relatively large amounts of water in this sub-humid to humid precipitation zone. Padre Island National Seashore, Texas. Hibernal aspect, February. FRES No. 39 (Prairie Ecosystem). K-66 or K-69 (a form of Bluestem Prairie): map units explained with two preceeding photographs. SRM 723 (Sea Oats). Beardgrass Series of Brown et al. (1998). Western Gulf Coastal Plains- Laguna Madre Barrier Islands and Coastal Marshes Ecoregion, 34i (Griffith et al., 2004).

184. Sea oats stabilizing sand dunes- This small dunes (and those barely visible in the background) were on a recent deflation plain where they were invaded by sea oats. This was an example of conditions and sites on which a climax species can also function as a colonizing species, at least locally when propagules are in close proximity to disturbance. The key role(s) of sea oats are what distinguish this as the sea oats prairie. Padre Island National Seashore, Texas. February. At local area scale this is a unique form of FRES No. 39 (Prairie Ecosystem), K-66 or K-69. SRM 723 (Sea Oats). Western Gulf Coastal Plains- Laguna Madre Barrier Islands and Coastal Marshes Ecoregion, 34i (Griffith et al., 2004).

185. Big bluestem on the sea oats-seacoast bluestem prairie- That the prairies and marshes of the Gulf Coast are the southern extremity of the general tallgrass prairie vegetational region, Kuchler's Bluestem (Andropogon, Panicum, Sorgastrum) Prairie, is illustrated by local dominance of big bluestem on a microsite in the tallgrass variant form referred to as sea oats prairie. This grassland developed on a barrier flat which is a more-or-less level area of land on a barrier island that formed between the foredunes and the back-island dunes. Barrier flats usually originated as wind-deflation flats. The deflation flats formed when wind erosion removed loose materials like sand. Generally, the wind erodes sand and/or earth down to the levels at which moisture from the surface water table holds the sand in place, the particles remaining being either too heavy or bound to each other to be picked up. The eroded materials are re-deposited as dunes which are drier than the moist sand remaining (un-eroded) just above the ground water level. Wind not only moves geologic materials directly but also indirectly as wind-driven tides which can come from both the sea (eg. the Gulf) and from any bodies of water situated between the barrier island and the mainland (eg. the Laguna Madre to the west, the mainland-side, of Padre Island). When the wind-driven tides from saline lagoons (eg. Laguna Madre) flood lower portions of islands this salty water carries away suspended soil and parent materials like sand (ie. water erosion). It also floods these low sites with water several times saltier than ocean water. These processes are the basis of the dynamic nature of dune landscapes such as often found on lake shores and barrier islands. Barier flats and local topographic variations within them often are more favorable for plant production. This portion populated by big bluestem may have been a more fertile or, as is generally the case for wind-deflation flats,a more mesic microsite. The size, habit, and physiogonomy of the big bluestem stand attest to the affinity of the general sea oats-seacoast or little bluestem prairie as a part of the tallgrass prairie. From the standpoint of plant succession the vegetation on these flat portions of barrier islands are the equivalent of the regional climax with the barrier flat being the peneplain in the geologic cycle perspective of William Morris Davis that was the basis of the Clementsian monoclimax theory. The occurrence of big bluestem as a local dominant and seacoast bluestem as overall dominant is evidence for the affiliation of this relatively restricted range type with the general bluestem prairie that extended over so much of the latitude of central North America. Padre Island National Seashore, Texas. hibernal aspect, February. FRES No. 39 (Prairie Ecosystem). K-66 or K-69 (one form of the Bluestem Prairie). SRM 723 (Sea Oats). Beardgrass Series of Brown et al. (1998). Western Gulf Coastal Plains- Laguna Madre Barrier Islands and Coastal Marshes Ecosystem, 34i (Griffith et al., 2004).

186. Composite view of the seacoast bluestem-sea oats prairie- This vegetated barrier flat supported a diverse plant community which served as a general model in minature of the coastal prairie and marsh portion of the tallgrass (= bluestem) prairie. The nature of a barrier flat in the interior of a barrier island and the general geologic processes forming barrier flats was presented under the immediately preceding slide. As explained above, barrier flats begin as wind-deflation flats (= wind-deflation troughs). Within the general wind-deflation trough or flat (which in time becomes a vegetated barrier flat) there are lower areas running across (crossways or perpendicular to) the general or whole barrier flat that extends roughly from the foredunes to the back-island dunes. These deeper but narrower wind-deflation troughs occur within the general deflation trough extending to various distances across it thereby creating a micro-relief that can be pictured as a washboard or corrogated sheet metal. These deeper wind-deflation flats ("mini-flats") formed during droughts or periods when the sand was drier than that for the average of the entire time sequence over which the whole, the overall, deflation flat formed. As the sand was drier for a greater depth in the soil profile the wind could erode away more of the profile (ie. "deeper down"). Over time the result was a micro-topography of troughs within the overall or general trough (ie. much like a washboarded road). When rainfall returned to the average for that of the entire chronological sequence under which the general or entire wind-deflation flat (and, ultimately, the barrier flat) formed the deeper "trenches" (ie. the narrower wind-deflation troughs) filled up with water. During periods of average or above average precipitation the ground water table is closer to the land surface of the barrier flat such that water rises to the surface in the deeper but narrower troughs. This creates local areas of sub-irrigated prairies, fresh water marshes, or small, often ephemeral, lakes within the extent of the overall barrier flat (the vegetated land located between the dunes at the seaward and mainland edges of the barrier island). Such conditions existed on the heavily vegetated barrier flat shown in this slide. A fresh water lake formed by ground water rising to the surface of one of the deeper, drought-induced, small wind-deflation troughs in the base floor of a wind-deflation-derived basal flat (far horizon). Subirrigation was the edaphic penomenon responsible for the wet (at least ephemerically wet) prairie or, perhaps, the marsh community in the foreground of this landscape. The round, conspicuously green leaves are marsh or largeleaf pennywort (Hydrocotyle bonariensis), an indicator species of very mesic to hydric soil conditions. Sandwiched between the lake and its adjacaent marsh and the wet prairie is the predominant mesic edaphic habitat supporting bluestem prairie the dominant of which was seacoast bluestem. The combination of natural fresh water lakes and abundant forage from tallgrass prairie and marsh made the landscape of this barrier island ideal habitat for numerous species of wildlife from waterfowl to white-tailed deer. With coming of the white man it served as the natural resource base for ranching empires. Research into the cattle-ranching frontier revealed unequivocally that the roots of this industry were in the soil of the coastal prairies and marshes area. (See for eg. Jordan, 1993). The barrier flats of such barrier islands as Padre Island served as critical cattle range for several ranching empires, including for a brief period King Ranch. Students can get a good introduction to the geology, ecology, and human history of barrier islands from the Padre Island guide book by Weise and Shite (1980) from which much of the above explanation was extracted. Padre Island National Seashore, Texas. Hibernal aspect, February. FRES No. 39 (Prairie Ecosystem). K-66 or K-69 (one variant of Bluestem Prairie). Inherent problems with large-scale mapping units such as the Kuchler units of potential vegetation were discussed with the first two slides that introduced this rangeland cover type (SRM 723, Sea Oats). Beardgrass Series of Brown et al. (1998). Western Gulf Coastal Plains- Laguna Madre Barrier Islands and Coastal Marshes Ecoregion, 34i (Griffith et al., 2004).

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