True Prairie - IA

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True prairie was the most nearly completely destroyed major unit of the grassland biome in North America. It was one of the easiest and first of the major grasslands in North America to be converted into farm fields. In contemporary times true prairie is the rarest, most restricted, and least known of the vegetational units that Clements (1920, ps.121-131) designated as associations of the grassland climax. In fact, true prairie was rare even when the first generation of vegetation scientists such as the great grassland ecologists like Frederic E. Clements and John E. Weaver (along with classmate Roscoe Pound) and their students like F. W. Albertson and G.W. Tomanek, the second generation of grasslanders, set out to describe North American vegetation.

Clements (1920, p. 121) described the true prairie, the Stipa-Koeleria association, in this way: "The true prairies occupy a distinct belt between the subclimax and mixed prairies, reaching from Manitoba to Oklahoma. This position as well as their relationship is shown by the presence of Andropogon scoparius derived from one and Stipa comata from the other". Stated more specifically, true prairie is a major grassland--a Clementsian association and, later, a cover type (or group of cover types)--between the bluestem-dominated tallgrass prairie, the subclimax prairie (Andropogon associes) in Clements' monoclimax model (Clements, 1920, p. 131), to the moister east and the needlegrass-dominated mixed prairie to the drier west.

Weaver and Clements (1938, ps. 519-520) described the true prairie as forming "a broad ecotone" between mixed prairie and the tallgrass prairie with invasion by tallgrass dominants, especially big bluestem (Andropogon gerardii) and Indiangrass (Sorgastrum nutans) into the original true prairie

Other than the general relative spatial position between tallgrass and mixed prairies little can be determined about true prairie when compared to the other major North American grasslands. "Cultivation has perhaps destroyed this association to a larger extent than other community of the grassland, and its limits are accordingly difficult to trace" (Clements, 1920, ps. 121-122). Gradual spatial changes in climate and affect of topographic (relief) variables along with "the all but complete removal of the original cover over large areas" resulted in the situation in which "the exact limits of the several prairies can never be set, and the boundary lines drawn on any map can be only general approximations" (Weaver and Clements, 1938, p. 519).

Less is known about species compostion, structure, resilence, successional patterns, soils, and so on for true prairie than for any major unit of the central grasslands. In fact, it is easier to locate and distinguish some minor climax grassland communities (eg. bamboo canebrakes) than the once widespread magnificant true prairie. One of the more interesting things about descriptions of true prairie were revisions made in as to dominant species. As was quoted above in Clements' original description of true prairie there was no mention of any Sporobolus species. Stipa spartea and S. comata from the mixed prairie and Andropogon scoparius from tallgrass prairie were defining dominants with Koeleria cristata and Agropyron glaucum (=A. smithii) forming the other consociations (Clements,1920, p. 121). Later in what was probably his most famous paper, Nature and Structure of the Climax, (Clements, 1936, p. 273) presented more of his famous (infamous to detractors) terms including eudominant(s), "one or more dominants peculiar to [the association]". Clements' example was eudominants of the true prairie association: Stipa spartea, Sprorbolus asper and S. heterolepis. This interpretation was subsequently included in the famous textbook, Plant Ecology, (Weaver and Clements, 1938, p. 518): "The major dominants are Stipa spartea, Sporobolus asper, S. heterolepis, Andropogon scoparius, Koeleria cristata, Agropyron smithii, and Bouteloua curtipendula, often with Andropogon furctus (=A. gerardii) and Sorgastrum nutans from the postclimax [tallgrass prairie] and Stipa comata from the mixed paraire. .. Stipa spartea, Sporobolus asper, and S. heterolepis are the three most characteristic dominants, since they do not occur as such in any other association". Along with porcupinegrass, tall and prairie dropseed were the defining dominants in the revised description of true prairie.

In the post-Clementsian period the distinction between true prairie and tallgrass prairie (Pound and Clements, 1900, ps. 348-350; Weaver and Clements, 1938, ps. 518-523) has been largely forgotten with the two major types or associations used synonymously or one for the other as, for instance, by Rissser et al., (1981). A more prominent example was that shown by Kuchler (1964, 1968) who did not describe or map true prairie or a unit dominated by needlegrasses, ,dropseeds, and little bluestem (a Stipa-Sporobolus-Andropogon unit). Strangely enough, however, noneother than John Weaver contributed to this confusion and apparently used true and tallgrass prairie more or less synonymously when he associated true prairie much more with big bluestem than with prairie dropseed (Weaver, 1943; Weaver and Bruner, 1954), but then he recognized little bluestem, needlegrass, and prairie dropseed as the dominant species of upland prairie or true prairie (Weaver, 1965, ps. 68-76)

In the treatment of true prairie in this chapter the current author followed Weaver and Clements (1938, p. 518) and Weaver (1965, ps. 68, 74) and used dominance by needlegrasses, little bluestem and, especially, prairie or tall dropseeds as the criterion is determining examples of true prairie. This was also more-or-less consistent with the subsequent treatment by Dodd (In: Gould, 1968, ps.325-328) as explained in the first photo-caption immediately below.

Historical note regarding mapping of natural vegetation and recognition of rangeland cover types: For whatever reasons, both Kuchler (1964, 1966) in mapping of the potential natural vegetation of North American and the Society for Range Management in naming and describing rangeland cover (dominance) types (Shiflet, 1994) failed to include true prairie! These were unexplained glaring omissions. In fact, these were remarkable oversights given the documentation, detailed descriptions (including distinctions among true prairie and tallgrass and mixed prairies), historic coverage, and general textbook recognition of this major unit (whether interpreted as type, association, or general community) of native vegetation.

Omission by the Society for Range Management from Rangeland Cover Types of the United States (Shiflet, 1994) was likely the result of having no one to write up a description of this dominance type. It was remarked in the Range Cover Types section of the Introduction herein that in Shiflet (1994) some regions and areas therein were treated with considerable detail (cover types had been identified, named, and described by vegetation "splitters") whereas other regions had gotten the proverbial "short shrift" for what appeared to be a simple explanation that either nobody was asked to submit a contribution or there was not anybody qualified or willing to do so. The "conspicuous by its absence" of true prairie (Stipa-Sporobolus-Andropogon-Koeleria) from the nearly comprehensive works of Kuchler (1964, 1966), who was a professor in, of all places (and note the irony of this), Kansas defined explanation- or even belief that such an erroneous omission could have been made. Absence of true prairie from Shiflet (1994), omission of a needlegrass-dropseed-bluestem-Junegrass rangeland cover type, was probably due in part to the fact that authors of Northern Great Plains Cover Types in Shiflet (1994, ps. 69-84) relied on the units of natural range vegetation recognized by Kuchler (1964, 1966), including the common (English) name titles, for their cover (= dominance) types. They simply repeated Kuchler's erroneous omission.

This chapter devoted to True Prairie, even if brief, was included to recognize this major and distinctive North American grassland even though it had been mostly destroyed by the time it was first recognized and described by Clements (1920, p. 121). And, no (in anticipation of the obvious question) that is not the reason why Kuchler (1964, 1966) omitted a unit entitled True Prairie, Needlegrass-Dropseed-Bluestem-Junegrass (Stipa-Sporobolus-Andropogon-Koeleria). He recognized the "extinct" California Steppe (Stipa) and that unit of potential natural vegetation was displaced permanently (in human time scale) by the naturalized unit, California annual grassland. Currently there are far more acres of Stipa-dominated true prairie than of Stipa-dominated California or Pacific bunchgrass prairie. No, Kuchler (1964, 1966) just missed it, plain and simple; and nobody else appears to have caught the error (or had the moxey to point out the mistake). The paultry effort herein was a step toward rectifying a major and "replicated" oversight. (Maybe this recognition will somewhat settle the ghosts of Drs. Weaver and Clements.)

1. Frederic E. Clements, arguably the greatest range ecologist of all time, and John E. Weaver, the man who know more about the North American prairie and the grasslands of the Great Plains than anyone who ever trod sod, were adamant that true prairie was the dominant climax form of prairie per se and not bluestem-Indiangrass tallgrass prairie, which was a separate form, another vegetation type in current parlance. True prairie in their view (Weaver and Clements, 1938, ps. 518-519) "occupies a fairy distinct belt between the tall-grass and mixed prairies" extending from southern Manitoba through central Kansas and into southern Oklahoma. However, "cultivation has almost clompetely removed the true prairie over most of its area" with natural boundaries having to be hypothetically reconstructed from small, scattered remnants. (Weaver and Clements, 1938, p. 518) listed major dominants of true prairie (in order): Stipa spartea, Sporobolus asper, S. heterolepis, Andropogon scoparius, Koleria cristata, Agropryon smithii, and Bouteloua curtipendula "often with Andropogon furcatus [= gerardii] and Sorgastrum nutans from the postclimax". Weaver and Clements in both editions [1929 and 1938] of Plant Ecology held to this interpretation as did their colleague Shelford (1963, p. 334) .As to tallgrass species, Weaver and Clements explained how removal of buffalo and decrease of fires led to replacement of even short grasses like buffalograss (Buchloe dactyloides) by the bluestems (Andropogon spp.) and other tallgrass genera. They noted " the belief of the pioneers that the bluestems (Andropogon) followed in the wake of the settlers and drove out the buffalo grass ". " The advance of the tall Andropogons over much of the true prairie is attested by scientific accounts of its composition" (Weaver and Clements, 1938, ps. 458-460, 516, 518-521).

Dodd (In: Gould, 1968, ps.325-328) provided a detailed description of true prairie, but he erred by omitting tallgrass prairie and, instead, including it in true prairie thereby eliminating a description of the tallgrass grassland, the Kuchler-66 (Bluestem Prairie), and therby overextending the range of true prairie eastward. Nonetheless, Dodd did recognize the Stipa-Sporobolus dominated climax grassland communities (ie. true prairie) as a major and geographically large, even immense, (regional scale) unit of potential natural vegetation. Dodd was unique among more recent (post-Clementsian era) workers in this interpretation, and he was correct. In describing rangeland cover types, the Society for Range Manageament (Shiflet, 1994) was remiss in leaving out the Stipa-Sporobolus (with associates like A. scoparius, K. cristata, and B. curtipendula) range type. Ditto for omission of the Clementsian true prairie in standard contemporary descriptions/discussions of North American grasslands. For example, Simms and Risser (in Barbour and Billings, 2000, ps. 333-345) excluded true paririe after previously listing (in Barbour and Billings, 2000, p. 331) the climax grasslands of Weaver and Clements (1929, 1938), including the Stipa-Sporobolus true prairie association. Obviously true prairie is no longer a major existing unit of North American grassland which is apparently the reason for its later exclusion by Sims and Risser, but it does exist and is a major range type to livestock operators, nature preservtionists, biologists, and other conservationists dependant on and concerned about its wise use and proper management. Furthermore, as a unit of potential natural vegetation-- a transition (" abroad ecogtone"[Weaver and Clements, 1938, p. 519]) between tallgrass and msxed prairies-- it should be recognized. Even Kuchler (1964, 1966) missed the Stipa-Sprorbolus climax grassland as potential natural vegetation.

Weaver and Clements (1938, p. 518) interpreted S. spartea, S. asper, and S. heterolepis as the "most characteristic dominants" because these species did not exist as dominants in any of the other climax grasslands (associations). To these pioneer plant ecologists who knew the grasslands at the time when they were being plowed under it was this feature of restricted dominance rather than extent or area of land dominated that was key and therefore indicative of true prairie, the ecotone between tallgrass and mixed prairie associations. Theirs' was a continent-wide perspective as the great grasslands were being highly altered-- in many cases destroyed-- by industrial man. It was a synoptic view that none thereafter could see. For this reason, current and future ecologists have no choice but to accept the eye-witness accounts of these pioneers of Grassland Ecology.

Here is a sample of true prairie, and one of the John Weaver's favorites. Scene of Nine-Mile Prairie (9 miles form his beloved University of Nebraska). The three characteristic dominants of true prairie were porcupinegrass (Stipa spartea), prairie dropseed (Sporobolus heterolepis) and tall dropseed . Besides these species this view includes little bluestem, Junegrass (Kolera cristata) and the introduced smooth bromegrass (Bromus inermis). August and, most appropriately, a drought. Lancaster County, Nebraska. FRES No. 39 (Prairie Ecosystem). No Kuchler or SRM designation (remarkable given the body of evidence that supports distinction between true and tallgrass prairies).

2. Sample of true prairie in Ozark Plateau- A relict of true prairie (actually an ecotone between tallgrass prairie and true prairie) on an upland prairie in the westrn Ozark Highlands (Springfield Plateau) dominated by porcupinegrass (Stipa spartea) with roundseed rosette panicgrass (Panicum sphaerocarpon, subgenus Dichanthelium) locally common and Ohio spiderwort (Tradescantia ohioensis) well-represented. There were also a few plants of Junegrass (Koleria cristata) and of the native forb, western yarrow (Achillea lanulosa= A.millefolium) in this vernal society.

Burkhart Prairie, Newton County, Missouri. Late May; peakstanding crop, ripening grain stage of phenology.

3. Sample of true prairie dominant- Local stand of porcupine grass on a native transtion grassland, an ecotone between true paririe and tallgrass prairie, in the western Ozark (Springfield) Plateau. There were also some plants of western yarrow; dewberry (Rubus flagellaris); and the naturalized invaders, tall fescue (Festuca arundinacea), a perennial agronomic species, and Japanese chess (Bromus japonicus), an annual Eurasian invader.

Burkhart Prairie, Newton County, Missouri. Late May; peakstanding crop, ripening grain stage of phenology.

4. Prickly grass- Examples of porcupinegrass in the spring society of a tallgrass prairie in the Prairie Peninsula at far-western edge of the Ozark Highlands (Springfield Plateau). The first photograph presented a local population or stand of porcupinegrass while the second photograph featured one individual plant of this cool-season, C3 grass. Stipa is the major genus of the Stipeae tribe. Recent taxonomic revisions (Barker et al., 2007, ps. 156-161) based on cladistic interpretations resulted in major changes in nomemclature (notably a proliferation of genera not previously recognized in North America) along with rearrangement--and consequent renaming--of species heretofore included in the genus Stipa.

Burkhart Prairie, Newton County, Missouri. Late May; peak standing crop, grain ripe/shedding stage.

5. Whole aboveground plants- Two plants of porcupinegrass (first slide) and details of upper shoots (complete with inflorescences) of these plants (second slide). These pants were growing in the virgin sod of a Ozark Plateau prairie that had been used as a prairie hay meadow for well over a century.

Burkhart Prairie, Newton County, Missouri. Late May; peak standing crop, ripening-grain stage.

6. Sexual shoot and sexual adornment- Two sexual shoots in one plant of porcupinegrass (first slide) and a single inflorescence of porcupinegrass (second slide) in a prairie hay meadow in the Springfield Plateau of southwestern Missouri. Prairie hay had been mmade off this nice meadow for over a century.

The inflorescence of the Stipa species is interpreted as a panicle.

Your author hauled hay (the rectangular bales that weighed roughly 65 to 70 pounds) off of this meadow when he was hillbilly of high school age. In those days even many of the dud high school students enjoyed hauling and barning hay believing that it got them in shape for football scrimmage, and of course football was what high school was about. Now no one wants to work so even prairie hay is put in big round bales (half the hay rots before the stock can eat it). Would this author do this all over again? You betcha! There is a fierce sense of accomplishment when hard work is done. This hilbilly still comes back to visit the procupinegrass. Hopefull it will still be there many human generations from now. (Don't bet on football dying out either, but one can always it will eventually be put in its proper perspective.)

Burkhart Prairie, Newton County, Missouri. Late May; peak standing crop, ripening-grain stage.

7. Panicles of quills - Inflorescence (panicle) of procupinegrass on a native prairie hay meadow in the western Ozark (Springfield) Plateau. Burkhart Prairie, Newton County, Missouri. June.

8. Five graminaceous quills- Five ripe and shed caryopses of porcupine grass showing entire fruit, including awns of lemmas (first slide), details of the tightly twisted lemmas (second slide), and grain portion, including sharp callus of lemmas (third slide).

Viewers can eaasily see why Stipa spartea is potentially one of the most mechanically injurious of all North Amereican range plant species. Enter one of the Four Cardinal Principles of Range Management: Proper Season of Use. Given that 1) porcupinegrass is a cool-season species and 2) that it can be mechanically harmful to grazing animals, it follows that ranges with substantial cover of porcupinegrass whould be grazed juniciously in early to mid spring before the sharp calluses and twisted awns of the lemmas develop and/or grazed properly again in mid-summer after grains have been shed and are not longer positioned to damage eyes, muzzles, delicate skin, etc.

Burkhart Prairie, Newton County, Missouri. Late May; grain ripe/shedding stage.

9. Bunched base- Basal shoots of a single cespitose plant of porcupinegrass at full-shoot (including inflorescence) stage of phenology. Virgin sod of a prairie in the far-western Ozark (Springfield) Plateau.

Burkhart Prairie, Newton County, Missouri. Late May; peak standing crop, ripening-grain stage.

10. Smooth shoot of a prickly top- Details of culm and leaves of porcupinegrass. Blade, sheath, and ligules (first slide) and relation of leaf axil to culm (second slide) in plants growing in the Prairie Penisula portion of the Ozark Plateau.

Burkhart Prairie, Newton County, Missouri. Late May; peak standing crop and full maturity.

Taxonomic reminder: Readers wishing nomenclature and systematics of traditional Stipa species, including S. spartea, as changed based on cladistic criteria were herein referred to Barkworth et al. (2007, ps. 156-161). Porcupinegrass (along with other phenetic Stipa species such as S. comata, needle-and-thread) was moved to a genus not recognized in North America by traditional taxonomic treatment.

11. Canada wildrye (Elymus canadensis)- This is a consociation (a community having a single dominant species, sometimes having a single species period as a natural single species stand; synecological term coined by F.E. Clements) of Canada or nodding wildrye on a floodplain on the Texas Grand Prairie (a tallgrass prairie). The wildryes (Elymus spp.) are one of the more important and widely distributed genera of cool-season grasses in the bluestem-dominated tallgrass prairies and oak-hickory-tallgrass savannas. They are not dominants of true other than at local scale, but they certainly do occur on and characterize the type on more mesic habitts. Erath County, Texas. June. FRES No. 39 (Prairie Ecosystem), one form or subunit of K-669 Bluestem Prairie).

12. Inflorescene (spike) of nodding Canada wildrye - Erath County, Texas. June.

13. Virginia wildrye (Elymus virginicus)- This wildrye species grows on the same general range types and geographic region as nodding or Canada wildrye, but typically on more mesic and shaded microsites. Frequently, however, these two species literally grow side-by-side, and according to Gould (1975, p. 167), readily hybridize. An instructive exercise for beginning Agrostology students is to use different manuals (eg. Hitchcoch and Chase, 1951 vs. Gould, 1975) and find that some specimens will key to E. canadensis using one authority and to E. virginicus using a different author depending, of course, on the number of Elymus species in the key. Both species are highly palatable decreasers extending from the eastern edge of tallgrass prairie through true prairie to mixed prairie. The vertical shoots of the example shown clearly designate it as E. virginicus. Border of tallgrass hay meadow in island of Cherokee Prairie in Ozark Plateau. Ottawa County, Oklahoma. June.

14. Individual plant of prairie dropseed- This is the most widespread dominant species that defines the true prairie range type. Only little bluestem and the two needlegrass species could vie with this species as a widespread dominant of true prairie. Shown here is the typical habit of this cespitose species. Scale is shown by the meter-long walking stick. Hay meadow, Ottawa County, Oklahoma, July.

15. True member of true prairie- Local population of conspicuously cespitose plants of prairie dropseed. This local consociation was growing in the heart of the tallgrass prairie. Nonetheless, prairie dropseed is a dominant of the true prairie to the immediate west of the tallgrass region. These plants presented the physiogonomy at close spatial scale of true prairie. Prairie dropseed is a (sometimes, the) defining species of true prairie. In the southern extremes of the true prairie prairie dropseed is commonly the most distinctive species defines this once widespread--and now nearly extinct--range type.

This "photoquadrant" was taken in the eastern Smoky Hills physiographic province.

Saline County, Kansas. Late June; pre-bloom phenology.

16. Two nice specimens of a dominant- Two plants of prairie dropseed on a tallgrass prairie in the wetern Ozark (Springfield) Plateau during moist times. The narrow, lacy, drooping leaves on a cespitose habitat are characteristic of this dominant bunchgrass. Throughout much of the Ozark Highlands of southwester Missouri prairie dropseed is found in association with the Four Horsemen of the Tallgrass prairie--big bluestem, little bluestem, Indiangrass, and switchgrass--obviously on climax or, at least, high seral stage grasslands.

In other words, even though these two plants were on tallgrass rather than true prairie they were typical climax grass species for this region and physiographic province.

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

17. The following series of slide-caption sets presented the habit (outward or general morphologic appearance) of prairie dropseed:

17A. Two examples- Two plants of the cespitose (tufted or bunched) prairie dropseed. This bunchgrass is the principal defining dominant of much of the true prairie. Most of the easily plowed bunchgrass prairie had already been turned under and converted to farmland by the time that Frederic E. Clements described it around the time of World War I (Clements, 1920, Weaver and Clements, 1929).

Newton County, Missouri. Early August; vegetative growth completed.

17B. Two more for good measure- Two more specimens of prairie dropseed were included because this former climax dominant of true prairie is now one of the most restricted of the original dominant grasses. Clements (1920, ps. 121-122) remarked that cultivated agriculture (field crop production) had likely "destroyed" true prairie to a relatively greater extent than other major grassland types ( Clementsian associations). In this earlier work Clements (11920, ps. 121-127) did not regard prairie dropseed as a dominant (or even a major) grass species of true prairie, but in later work (Weaver and Clements, 1938, p. 518) porcupinegrass, prairie dropseed, and tall dropseed were interpreted as "... the three most characteristic dominants [of true prairie], since they do not occur as in any other association". These three are cespitose species (bunchgrasses) in contrast to sod-forming grasses like western wheatgrass (Agropyron smithii) or intermediate species like little bluestem and sideoats grama (Weaver and Clements, 1938, p. 518).

In personal communications with "old-timers" such as farmers who knew pioneer (sod-breaking) farmers, your author ascertained that it was considerably easier to convert the bunchgrass form of true prairie to farm fields in contrast to the sod form of true prairie (it required considerably less effort to break the virgin sod of the tufted grass type with its more open and widely spaced perennial grasses). For this reason (and perhaps others), the native cespitose grasses like prairie dropseed, tall dropseed, and porcupinegrass were some of the first grassland plants to be reduced--often essentially estirpated--by earlier stages of prairie (former prairie) settlement.

Regardless, your well-traveled prairieman author has had more difficulty in locating prairie dropseed and porcupinegrass than any other plant species that was a climax grassland dominant. Hence, he felt oblidged to share "plenty" of images with his students. Enjoy: these blasted things are hard to find.

Specimens of prairie dropseed shown in these two images as well as those in the immediately succeeding photographs provided the typical open crown or "sprawling" habit of this cespitose species.

Newton County, Missouri. Late July; vegetative growth completed, immediate pre-bloom phenology.

17C. The cespitose habit- Several palnts (first slide) and a single plant of prairie dropseed on a small chert glade in the Springfield Plateau. Behind the single prairie dropseed plant in the second slide was a plant of prairie dock (Silphium terebinthinaceum). These two species are typically associated with one another.

Prairie dropseed is typically more characteristic and indicative of true than tallgrass prairie, but this species is found on grasslands of both of these Clementsian associations-- when they are climax and higher seral stages (eg. subclimax).

McDonald County, Missouri. July; pre-bloom phenological stage.

18. The next series of slides presented the entire shoot complex and panicle inflorescences of prairie dropseed.

18A. At full development- One plant of prairie dropseed at full bloom (right side, upper slide) and an overall view of its panicle inflorescence (lower slide)Range plant to immediate left of prairie dropseed in the first of these three slides was

Newton County, Missouri. Late July; peak bloom and peak standing crop.

18B. Panicles aplenty-Two panicles of prairie dropseed in early, still largely unexpanded, stage (first slide) and then in late anthesis with secondary branches of panicles fully expanded or extended (second slide).

Newton County, Missouri. Late July; peak bloom and peak standing crop.

18C. Co-dominant pals- Inflorescences of prairie dropseed and sideoats grama, two species generally regarded as mid-grasses of true prairie.

Newton County, Missouri. Late July; peak standing crop and anthesis stage of phenology.

Newton County, Missouri. Late July.

19. Two from the side- Side views of two fine specimens of prairie dropseed, the defining dominant species of much of the true prairie, almost all of which had been destroyed by the turn of the Twentieth Century (Clements, 1920; Weaver and Clements, 1929). In fact, so little of the true prairie remains that even many, if not most, grassland ecologists, range management specialists, wildlife biologists, etc. incorrectly/improperly use true prairie and tallgrass prairie as synomymous. For example, this error was illlustrated by the title The True Prairie Ecosystem (Risser et al., 1982) which was the synopsis work of the International Biological Program for the tallgrass prairie ecosystem.

Dominance of grassland, the prairie, vegetation by prairie dropseed, more than any other plant species, readily distinguishes between these two major climax grassland communities (associations or sub-units of the North American biome).

Newton County, Missouri. Early August; vegetative growth completed.

20. At its base- Basal view of a prairie dropseed plant at maximum shoot development, peak standing crop.

Newton County, Missouri. Early August; vegetative growth completed.

21. Characteristic leaves- Leaves of prairie dropseed at their maximum size and development (peak standing drop of the annual herbage growth-production cycle). Mature leaves of prairie dropseed taper to a fine point that is round in cross-section. This darning needle-like feature is a key distinguishing vegettive characteristic of this dominant of the true prairie.

Newton County, Missouri. Early August.

22. In the green- Fully expanded, full-grown panicle of prairie dropseed in early stages of fruit development and maturation.

Newton County, Missouri. Early August; fruit at milk stage of phenology.

Green around- Young female giant walkingstick (Megaphasma denticrus) on shoot of prairie dropseed. The author could not find any sources that indicted whether or not the giant walkingstick could change colors as a form of camouflage. Individuals of this species are of various colors with some walkingsticks, like the female seen here, of such color as to be ideally camouflaged.

One thing is certain: giant walkingsticks are phytophagous (plant-eating; herbiverous) insects. This no need-to-be-green-with-envy specimen was seemingly enjoying a meal of prairie dropseed until a photographer temporarily interrupted her meal.

Newton County, Missouri. Late July.

23. Not frost (it was Epson), but winter- Epson Perfection 600 struck again and converted a fine slide of the dead herbage of prairie dropseed at winter dormancy into a "sparkledy" likeness that appeared to be bearing a heavy coating of frost. There was no frozen moisture on this herbage. Instead, this was a failed attempt to get two nice Provia 100F slides (that captured the typical habit and physical appearance of prairie dropseed plants in their winter palege) before viewers watching from Internet land.

Maybe loyal digital range inspectors got something of the idea inspite of Epson's "perfection".

Newton County, Missouri. Late December; dormancy stage of phenology.

24. Closer views, but still Epson-frosted- Dead herbage (necromass) of the tillers of a dormant specimen of prairie dropseed (first slide) and upper sportions of sexual shoots with panicles (second slide).

Newton County, Missouri. Mid-December; overwintering (plant dormancy) and unshattered ripe-fruit stage.

25. Bleached more by Epson (than sun)- yep, Epson's "perfection" in scanning struck again, and here again maybe faithful digital rangemen got a general idea of appearances of prairie dropseed panicles in their hibernal aspect.

Newton County, Missouri. Late December; dormancy stage of phenology.

26. Try two backgrounds- These are better views of prairie dropseed panicles. Upper photograph is obviously two inflorescence lying atop snow whereas the lower slide is of two flower clusters laid over the dead herbage of prairie dropseed in dormancy.

Newton County, Missouri. Late December; overwintering ripe-fruit stage.

27. The following series of images showed the panicle type inflorescence and achenes of prairie dropseed in the dormant grass shoot (tiller) and fruit-ripe phenological stage(s)

27A. Tops of tops- Upper portions of two panicles of prairie dropseed with achenes at fruit-ripe stage on their dead inflorescences.

Newton County, Missouri. Mid- December; dead standing crop (necormass) of dormant plant, but with viable achenes at the unshattered ripe-fruit stage.

27B. Achenes on the stalk- Portions of two panicles of prairie dropseed still containing fully matured and unshed achenes. Strictly speaking the fruit of Sporobolus species is an achene (compare achene and caryopsis in Shaw, 2012, ps. 1045, 1046) rather than a caryopsis, the fruit of almost all members of the Gramineae. This is so because the testa, seed coat, is not fused to the pericarp, ovary wall, and instead the seed is loose within the fruit (Hitchcock and Chase, 1950, p. 413; Gould, 1975, p. 286; Gould and Shaw, 1983, p. 282).

Newton County, Missouri. Late December; overwintering and unshattered ripe-fruit stage.

28. Very distinctive- Spherical achenes of prairie dropseed still attached via their pedicels to branches of panicles. The achenes of prairie dropseed are among the most distinctive fruits of all North American grasses. They are not only globose (globe-shaped), but also have a tiny, string-like, vermiform (worm-like) appendage on the pericarp (See Hitchcock and Chase, 1950, Fig. 601, p. 422; Gould, 1975, Fig. 163, p. 310).

Newton County, Missouri. Late December; overwintering and unshattered ripe-fruit stage.

29. Co-dominants in winter- Prairie dropseed (left) and little bluestem (right) on true prairie at the edge or local ecotone of chert glades and oak (Quercus spp.)-hickory (Carya spp.) forest in the Ozark (Springfield) Plateau. A light covering of snow accentuated the ungrazed, dead herbage of these two co-dominant warm-season, decreaser grasses. On climax true prairie little bluestem rather than big bluestem, Indiangrass, or switchgrass is the dominant panicoid grass. Whereas the defining cool-season dominant of climax true prairie is porcupine grass (see again above) the defining dominant warm-season species of climax true prairie is prairie and/or tall dropseed (see immediately below) often even more than little bluestem.

Newton County, Missouri. Early January; dormancy stage of phenology.

30. Kissing cousins- Tall dropseed (left) and prairie dropseed (right) are alternatively dominant or co-dominant warm-season dominants of climax true prairie. These two eragrostoid species often (even typically) have greater cover and biomass than the panicoid little bluestem on true prairie. No, at this distance (size of image subject), and even with assistance of snow to accentuate plant herbage, these two species are not distinguishable (or even discernable). Given limitations of Internet viewing, however, your range professor--within confines of the digital world--did his best from this Springfield Plateau ecotonal prairie to show joint or dual dominance of these two growing-side-by-side kissing cousin species.

The next slide-caption episode may have helped eye-straining students.

Newton County, Missouri. Early January; dormancy stage and ripe fruit stages of phenology.

31. Tops of kissing cousins- Sexual shoots with panicles of prairie dropseed (at left and right-leaning; behind) and tall dropseed (at right and left-leaning; in front) that grew in true prairie that developed on a local ecotone of chert glades and oak-hickory forest in the Ozark (Springfield) Plateau. Yes, these shoots occurred naturally in the positions photographed from plants that grew cheek-by-jowl on this patch of God's own lawn. These as-they-were images demonstrated to students the natural close proximity at which these co-dominant decreaser often grow on true prairie.

God's gift of a classic textbook example. Learn and enjoy.

Newton County, Missouri. Early January; dormancy stage and ripe fruit stages of phenology.

31. Tall (= meadow) dropseed (Sporobolus asper)- This species is actually a taxonomic "complex" with three or four (or more) subspecies or varieties plus perhaps other closely related Sporobolus species that are difficult to differentiate and which often hybridize. The specimen shown here in the imediately succeeding slide was strictly speaking meadow dropseed (Sporobolus asper var. drummondii= S. compositus var drummondii) the taxonomic variety found in the Grand and Fort Worth Prairies and Cross Timbers of northcentral Texsas (Diggs et al., 1999, p.1327). S. asper is commonly second only to S. heterolepis as a dominant or associate species on tallgrass and true prairie range types. Sporobolus is one of the few Gramineae genera whose fruit is an achene and not a caryopsis. Erath County, Texas. October.

32. Inflorescence of tall dropseed or, more specifically, meadow dropseed- The contracted panicles of this species complex are partially to completely enclosed within the surrounding leaf sheath (ie. at least some of the infloresences on individual tall dropseed plants remain within the boot from which fruits are shed with the eventual disintegration of this enveloping lower portion of the leaf). Erath County, Texas. October.

Sporobolus complex- A group of closely Sporobolus species, varieties of one species, or varieties of two or more species with a wide biological range have long been recognized as important on tallgrass prairie. Tall dropseed, as a catchall common name, has been treated as including up to three varieties of the species S. asper or, as a synonym, S. compositus. . The specific epithet asper means "tall". More recently it was discovered that the author of S. asper used as basis of his scientific name an earlier designation that by international botanical rules was "illegitimate" (Tyrl et al., 2008, p. 173) so that the binomial now regarded as correct is S. compositus.

If this confusion was not enough there has been the long-standing view that S. compositus (= S. asper) is so "morphologically variable" that perhaps it is "conspecific" with S. clandestinus (Tyrl et al., 2008, p. 173). All told this mess amounts to what taxonomists regard as a "complex". The Sporobolus compositus complex is a difficult assemblage of forms, perhaps affected by their primarily autogamous breeding" which is largely responsible for these plants being able to "perpetuate unique character conbinations" (Barkworth et al., 2003, ps. 121-122 ). Autogamy refers to self-fertilization within a flower. Much of this autogamy occurs in cleistogamous spikelets, those having "self-contained" fertilization (Gould, 1975, ps. 303, 631).

By whatever name(s)--at species or variety level--there are at least three taxonomic forms under the scientific "handle" for a generic tall dropseed. Presented immediately below were two of these: 1) "regular" tall dropseed (S. compositus var. compositus= S. asper var. asper) and 2) meadow dropseed (S. compositus var. drummondii = S. asper var. drummondii).

Weaver and Clements (1938. p. 520) regarded tall dropseed as a midgrass and while a dominant of true prairie a frequent member of tallgrass prairie. On tallgrass prairie tall dropseed commonly achieves heights of four to five feet, a stature roughly equal to that of little bluestem growing on tallgrass prairie Given this fact, it appeared to the current author that tall dropseed could be either a tall- or midgrass species depending on range site, especially edaphic features of it range environment.

Successional status of tall dropseed, especially Sporobolus asper var. asper (= S. compositus var compositus), is a matter of confusion to those who do not recgnize (ie. understand) the difference/distinction between tallgrass prairie and true prairie. Tall dropseed, again especially S. asper var. asper, can in some areas be a "Fifth horeeman of the Prairies" (behind big and little blusetsms, Indiangrass, and switchgrass). In this tallgrass guild tall dropseed is the moxt xeric (or least mesic) of the five species falling in right behind or, on some sites, just ahead of little bluestem in adaptation to drier soil conditions. Tall dropseed has an ecological niche very similar to that of little bluestem.

For instance, Steyermark (1963, p. 165)--a taxonomist more than an ecologist or vegetation scientist--specified that in Missouri the various taxonomic varieties of tall dropseed were "drought-resistant" and "... spread into overgrazed areas where bluestem (Andropogon scoparius) is prevalent". Most likely Steyermark (1963, p. 165) was describing true prairie where tall dropseed and little bluestem are often climax co-dominants and not overgrazed tallgrass prairie where big bluestem, Indiangrass, and switchgrass had been replaced by tall dropseed and little bluestem.These later two species were specifically listed as major dominants of true prairie by Weaver and Clements (1938, p. 518). Steyermark (1963, p. 165) was, however, "right on target" (as he almost invariably was) though he got it "backassards" when he specified "overgrazed areas". Weaver and Clements (1938, ps. 49, 458, 469) explained that tallgrasses, especially Andorpogon species, replaced true prairie (and even shortgrass) dominants with settlement by white man (at least initially after extermination of North American buffalo and before the plow eliminated true prairie). Heavy grazing by native herbivores, most notably bison and prairie dog, apparently consiituted a zootic climax (a Clementsian disturbance climax) in which herbivory benefitted either shortgrass and/or tallgrass species over midgrass species.

Whatever the cause-effect relations, tall dropseed is a climax dominant (tallgrass or midgrass depending on one's interpretations of shoot height) of true prairie.

33. Autumn attire- Stand (first, second, and third slide) and single cespitose plant (fourth slide) of tall dropseed in its late autumn state of foliage under dry, mild atmospheric conditions following a comparatively moist late (late summer-early autumn) growing season. These plants were growing in an outside fencerow on a mesic tallgrass prairie in the western Ozark Plateau.

Tall dropseed was regarded by Weaver and Clements (1938, ps 518, 520) as a mid-grass dominant of true prairie. These pioneer plant ecologists, the perfection of grassland ecologists, listed tall dropseed as second only to neeedle-and-thread as a dominant of true prairie (Weaver and Clements, 1938, p. 518). Tall dropseed is also locally commonly on tallgrass prairie as was the case in this mesic tallgrass community that was dominated by big bluestem (Andropogon gerardii= A. furcatus).

Ottawa County, Oklahoma. Mid-December.

34. Staying inside their shoots- Three progressively closer views of upper portions of sexual shoots of tall dropseed growing in an outside fencerow on a mesic tallgrass prairie iin the western Ozark (Springfield) Plateau. The contracted panicles of this tallgrass species characteristically remain partly to almost completely enclosed within the boot much like the more common and widely distributed sand dropseed (Sporobolus cryptandrus), a characteristic species of mixed prairie. Details of the enclosed contracted panicle of tall dropseed were featured immediately below.

Ottawa County, Oklahoma. Mid-December.

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

Ottawa County, Oklahoma.

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

First slide: Ottawa County, Oklahoma. Mid-December. Second slide: Erath County, Texas, Early November.

37. Still in the package- Contracted panicles of tall dropseed still in the dead boot (the surrounding or encasing leaf sheath and, perhaps, portion of leaf blade) which in this case was the terminal leaf of this sexual shoot. Shoots of thecespitose Sporobolus species are tillers.

Ottawa County, Oklahoma. Late Decenber;

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

Erath County, Texas, Early November, fruit-shatter stage.

39. Close-up of sward of true prairie- A consociation (to use Clement's term) of prairie dropseed but with its associate, prairie dock (Silphium terebinthinaceum), existing almost as a co-dominant.A bald knob, a dolomite glade (McClurg Glade), in the Ozark Mountains of Missouri. Ava Ranger District, Mark Twain National Forest, Ozark County, Missouri.Note the young sprouts of yellow wood or smoke tree (Cotinus obovatus).Maintained by prescribed burning. FRES No. 39 (Prairie Ecosystem). Not described by Kuchler, SRM, or Brown et al, 1998).

40. On the old Chisholm Trail, and a rare relict- Remnant climax range vegetation of true prairie co-dominated by little bluestem and prairie dropseed with big bluestem and Indiangrass as important tallgrasses, Canada wildrye as the major cool-season grass, and buffalograss (Buchloe dactyloides) as the most common shortgrass species. Sideoats grama (Bouteloua curtipendula) was an important warm-season grass .There was some Japanese chess or brome (Bromus japonicus), a naturalized Eurasian annual. The major forbs were pitcher or blue sage (Salvia azurea) and leadplant. There were scattered plants of skunkbush sumac (Rhus trilobata= R. aromatica), occasional thickets of Chickasaw plum (Prunus angustifolia), and a few well-dispersed American or white and slippery or red elm (few individuals of Ulmus americana and U. rubra, both of which grow throughout this area). Presence of elm as woody invaders likely was evidence of inadequate prairie fires (ie. an unnatural fire regime). This appeared to be self-evident given small size of these saplings. There were also some small groves with very large shoots of smooth sumac (Rhus glabra) growing on steep banks along outer margins of what had once been part of the Chisholm Trail, one of the most historically pivotical transportation routes of this part of the Cattle Kingdom.

This climax range plant community had developed on the south part of Wellington and McPherson Lowlands (formerly treated as part of the Great Bend Lowland, Great Bend Prairie, or Arkansas River Lowlands (Fenneman, 1931, ps. 27-28; Fenneman, 1938, ps. 616, 621; Frye and Swineford, 1949, ps. 71, 78-79; Schoewe, 1949, ps. 276, 292,-300 passim, respectively) portion of the Osage Plains (= Section) of Central Lowlands physiographic province or, as an alternative interpretation, in the Plains Border physiographic section (Frye and Swineford, 1949, ps. 71, 78-79). By any of these various perspectives this rangeland area was to the west of the Flint Hills. This relict tract of true prairie in this part of the subhumid zone was a "blend" of the more mesic tallgrass prairie to the humid east and the less mesic mixed prairie to the semiarid west. This was higher elevation rangeland just west of and adjacent to the Arkansas River Lowlands province by currently accepted physiographic maps

. Little bluestem was the dominant tallgrass species. The rest of the tallgrass element appeared to be represented more by the long-shoot Indiangrass (largest, tallest bunchgrasses in center midground of second photograph) than the short-shoot big bluestem. Prairie dropseed was the other dominant. Co-dominance by prairie dropseed and little bluestem a key diagnostic feature that defined this as true and not tallgrass prairie. Scarcity (quite limited cover and biomass) of both mid- and shortgrass species distiinguished this from mixed prairie.

This range vegetation was on a section of the historic Chisholm Trail that has been preserved just outside of Caldwell, Kansas, the so-called "Border Queen" that was just inside the Jawhawker State and beyond Indian territory which was one of several cow towns that served at one time or another as the terminus of this most famous of all the Texas cattle trails (Gard, 1954, ps. vii, 117, 153, 190, 197, 212, 236, 251, 259; Worchester, 1980, ps. xviii-xix, 130). Over a span of less than 20 years millions of head of Texas cattle were trailed over this fabled overland trade route. The romance, traditions, and virtues established on the Chisholm and the other fabled stock trails (including some for sheep and even hogs) are with us still, and by God's grace they will remain. One could not stand on this sod, gaze across the rangeland, and be unmoved. Our patriarchs toiled--even died--here and sanctified our profession and way of life. Holy ground to "hired men on horseback".

Sumner County, Kansas. June; late vernal aspect. FRES No. 39 (Prairie Ecosystem). Not described by Kuchler, SRM, or Brown et al, 1998). Central Great Plains- Wellington-McPherson Lowland Ecoregion, 27d (Chapman et al., 2001).

41. Sward of true prairie, and sacred ground- Relict tract of true prairie co-dominated by little bluestem and prairie dropseed on a preserved section of the Chisholm Trail. Indiangrass and big bluestem were the other tallgrass (and local associate) species.Canada wildrye was the main cool-season decreaser grass. There was some sideoats grama representing the warm-season midgrass component, but it was not common. Buffalograss was the principal shortgrass. The Eurasian cool-season annual grass, Japanese brome or chess, was locally abundant (on microsites). Most common forbs were blue or pitcher sage and leadplant. Barely visible in background of this "photo-quadrant" were scattered saplings of American and slippery elm, thickets (colonies) of Chickasaw plum, and some skunkbush sumac.

This was a closer-in view of the true prairie range presented in the immediately preceding two photographs. It was obviously Excellent range condition class. Current season grazing had been limited to wildlife and insects. (The photogrpher did glimse ghosts of longhorns and trail drivers, but reverently refrained from putting them on Kodachrome.) A trailing procession, including buffalo and the redmen of the Plains Tribes, had long blessed this sacred earth. View with adoration.

42. So-so specimen of true prairie- A range of true prairie in high Fair to low Good condition class in the subhumid zone. This grassland was obviously an ecotone, a transition zone, between tallgrass prairie to the east (in the Flint Hills) and mixed prairie to the west (the Redlands, Rolling Redlands, Rolling Red Hills, or Rolling Red Plains portion of the Great Plains). This range was in the Great Bend Lowland or Great Bend Prairie portion of the Plains Border physiographic section, itself a transition zone in the physiographic and geologic sense. Dominants of this cover (dominance) type unambiguously delineated this as part of the once extensive and now nearly obliterated true prairie described by Weaver and Clements (1938, ps. 519-520). Little bluestem and prairie dropseed "swpped places" as dominants of this grassland. There were a few plants of Indiangrass and Canada wildrye was well-represented, but purple threeawn (Aristida purpurea), sideoats grama (Bouteloua grama), hairy grama (B. hirsuta), and silver bluestem (Andropogon saccharoides) were far more abundant and comprised greater proportions (more cover and biomass) of the plant community on this range than on the relict tract on the Chisholm Trail just described and on the pristine range described below.

Grant County, Oklahoma. Late June; late vernal aspect. FRES No. 39 (Prairie Ecosystem). Not described by Kuchler, SRM, or Brown et al, 1998). Central Great Plains- Prairie Tableland Ecoregion, 27d (Woods et all., 2005).

43. True prairie on the Great Bend Prairie- True prairie (little bluestem and prairie dropseed-dominateed subhumid prairie) in the physiographic unit designated variously as Great Bend Lowland, Arkansas River Lowlands (including Great Bend Lowland, Wellington Lowland, and McPherson Lowland), and Great Bend Prairie (Fenneman, 1931, ps. 27-28 and Fenneman, 1938, ps. 616, 621; Frye and Swineford, 1949, ps. 71, 78-79; Schoewe, 1949, ps. 276, 292,-300 passim, respectively) Osage Plains (= Section) of Central Lowlands.

Two photographs presented two different views (vantage points) of the same range that was introduced in the preceding photograph. This range was in high Fair to perhaps low Good range condition class. These two slides showed the much greater botanical/floristic diversity (differences in species composition, physiogonomy, community structure, etc.) in the range vegetation on this degraded pasture (deteriorated range) as compared to the much less sprecies-rich and more consistent physiogonomy of the climax vegetation on a nearby (half mile distant) range that was in pristine condition (presented and described immediately below). The range vegetation shown here also departed substantially from that of the relict climax true prairie on the Chisholm Trail section described immediately above. The range plant community(ies) on this degraded range varied from dominance by species that were invaders and increasers (eg. purple threeawn, silver bluestem, and sideoats grama) to decreasers (including the overall dominants, little bluestem and prairie dropseed, relatively widespread Canada or nodding wildrye, and occasionally present Indiangrass). There were only a few forbs. Dotted gayfeather (Liatris punctata) was the most common of these. No woody plants were present.

Ecology lesson (basic concept in Vegetation Science): Viewers should note the drastic differences in species composition and plant community structure at local spatial scale between greater cover, density, etc. of the dominant climax grasses (prairie drpseed and little bluestem) in the first photograph versus the relative amount (cover) of bare soil and dominance of the local plant community by purple threeawn, silver bluestem, and sideoats grama in the second photograph. A mound of bare red soil (cattle played on it) was in the upper right corner for perspective and proof of the dramatic differences in species composition, plant cover, etc. at local (microsite) scale. Such differences (diversity, if that term is preferred) in range vegetation were not present at such short distances on either the relict tract of true prairie presented immediately above or the "mint condition" true prairie only a half mile away described in the immediately following photographs. These three examples had different combinations of range sites, but at the scale of range types they were comparable enough to show that certain measures of diversity such as number of species and plant life forms are not necessarily ecologically (successionally) desirable features of range ecosystems and landscapes. Climax vegetation is often less diverse (perhaps more boring to those wishing to botanize) than seral vegetation (ie. range in Excellent condition class may be less species-rich than range in states of retrogression). That was generally the case for true prairie in the area represented here.

Management lesson (application to range analysis and grazing practice): This was a degraded range not a depleted or exhausted range. The range vegetation on this pasture was unquestionably in a stage of retrogression that had departed from climax. The range plant communities (or overall community) on this true prairie sere corresponded to a higher rather than a lower seral stage. It was still dominated by the two climax (potential natural) dominant species (little bluestem and prairie dropseed), but on many microsites the dominant (often the exclusive) species were invaders or at best increasers. There were also some (infrequent) local habitats dominated by Indiangrass, a long-shoot, tallgrass species. Range condition class (high Fair to low Good) reflected this departure from climax (potential natural) vegetation, terminus of the sere, yet at a more successionally advanced seral stage. By judicious grazing management (adjustment of stocking rate, modification of season and/or duration of grazing use, manipulation of frequency and intensity of grazing) this range could be improved (ie. brought back to climax or at least to a higher seral stage--a successional status,--than it is now). This range (vegetation on this sere) could be improved to the successional stage that corresponded to obviously being in Good (perhaps even Excellent) range condition class. Such range improvement would ultimately correspond to greater grazing (carrying) capacity and higher--not lower--stocking rates. To achieve this range improvement might well require an initial reduction in stocking rate, deferment (nonuse= no livestock grazing) during the warm-growing season of little bluestem and prairie dropseed, the climax dominant range plants, shorter though more frequent grazing periods, and simply more careful attention to cattle (the kind of livestock using this range) grazing of the range herbage.

44. A rare find of true prairie- Classic example of true pairie with range plant communities alternating in dominance by either little bluestem or prairie dropseed or co-dominance by both of these climax decreaser species. The upland ecotypic form of switchgrass and big bluestem were the associate species. There was some presence of midgrasses including sideoats grama and silver bluestem, but these verged on trace amounts with the overwhelming composition based on estimated cover and biomsas being little bluestem and prairie dropseed (and "backed-up" by switchgrass and big bluestem). There were no forb species worthy of note. No trees or shrubs were present. There were some traces of Japanese chess ( a Eurasian, cool-season, annual grass) at edges of cow paths.

In both of these photographs prairie dropseed-dominated climax vegetation (Sporobolus hetrolepis consociation) was in the foreground and little bluestem-dominated range plant communities (Andropogon scoparius consociation) was in the background. Switchgrass, one of two associate species, was conspicuous in foregrounds of both "photo-plots". An example of vegetation on this range where the climax plant community was co-dominated by little bluestem and prairie dropseed (rather than a patchwork of the two consociations as shown here) was presented immediately below.

This jewel of climax true prairie was in virgin status. This range plant community was in "mint condition" even under cattle grazing. All-in all a superb specimen of relict natural vegetation being wisely used.

45. Textbook true prairie- A true prairie range in pristine state that developed on the Wellington and McPherson Lowlands physiographic province (as designated in current maps such as those by Kansas Department of Transportation) immediately to the west of the Flint Hills province. Another view of the true prairie rnge introduced in the preceding two photographs. This climax range vegetation was co-dominated by little bluestem and prairie dropseed, a combination that a "dead-ringer" for true prairie (versus tallgrass prairie). Big bluestem and the upland form os switchgrass were the associate species. There were trace proportions of sideoats grama, silver bluestem, and purple threeawn but essentially there was almost no midgrass or shortgrass component to this climax grassland community. Also, there were almost no forbs; nor were any woody plants present. (It would have been a "monotous" floristic mixture had it not been such a premier, and just downright beautiful, example of one of the rarest grassland cover types.)

It was obvious that--as first described by Weaver and Clements (1938, ps. 519-520)--this climax vegetation was an ecotone between tallgrass prairie and mixed prairie range types. It was also in the Plains Border physiographic section (Frye and Swineford, 1949) and the Wellington and McPherson Lowlands or, also, Great Bend Lowland (Fenneman, 1931, ps. 27-28; Fenneman, 1938, ps. 616, 621), Great Bend Prairie (Frye and Swineford, 1949, ps. 71, 78-79), or Arkansas River Lowlands (Schoewe, 1949, ps. 276, 291-300) at a blending or merger of the Central Lowlands and Great Plains physiographic provinces so as to represent and encompass even more of a transition zone (floristically, physiogonomically, physiographically, geologically).

Note: the examples of true prairie in Sumner County, Kansas and Grant County, Oklahoma were all upland prairies within the Great Bend or Arkansas River Lowlands. Fenneman (1938, p. 616) explained that much of the topography in the Great Bend section was upland and that it was at the scale of physiographic units (sections or provinces) that the general term of lowland applied.

The Holy Grain of prairiemen- In the section immediately below three examples (three separate ranges) of true prairie were presented. The first of these examples was the ultimate expression or development of what is now one of the rarest of major associations of North American grasslands. This range was being grazed by cow-calf pairs. The second example was a range that had not been grazed for several years so that much of the standing crop was dead herbage from the previous year. Taken together these two example ranges--two treasured relicts--presented physiogonomy, structure, and species composition of true prairie, a climax association that was once a broad zone of transition grassland situated between the tallgrass prairie to the wetter east and mixed prairie to drier west. First though, a clarification and a belated lesson for some ecological authors.

A remedial lesson in formations (climaxes) of central grasslands North America: Soil/Natural Resource Conservation Service range site descriptions of this range vegetation (Loamy Uplands range site) completely missed the mark. Descriptions read as this should be tallgrass prairie dominated by big bluestem (up to 30% ) while tall dropseed was minor (at 5%) (Norton County, Kansas Soil Survey, Soil Conservation Service, and Web Soil Survey, Natural Resources Conservation Service). Totally wrong: this grassland was textbook Clementsian true prairie from perspective of co-domination by little bluestem and tall dropseed down to Psoralea argohyllum and Erigeron modestus as major forbs. See Clements (1920, ps.120- 131) in Plant Indicators and Weaver and Clements (1938, ps. 518-520) in Plant Ecology. Many of the range (and , later, ecological) site descriptions of grasslands in eastern portions of the semiarid region of Kansas were written to imply that the climax range vegetation should be tallgrass prairie like that in the humid and subhumid Flint Hills and Osage Questas rather than true prairie along eastern margins of he semiarid Southern Great Plains. Such descriptions are incorrect and woefully misleading,. They reflect the fact that writers of such descriptions were unaware of the true prairie as the wide ecotone between tallgrass prairie and mixed prairie (Weaver and Clements (1938, p. 519) as was described and illustrated below.

Even some academics who should have know better perpetuated what has become an unfortunate error in describing and mapping North American grasslands. For example, the International Biological Program sythesis volume for tallgrass prairie was mistitled as The True Prairie Ecosystem (Risser et al, 1981). Confession time: this author himself out of ignorance once made the mistake to indicate that true and tallgrass prairie were synonymous. This error was on a tour of Nebraska prairies during the North American Prairie Conference. A more knowledgable (and much older) prairieman schooled under the immortal F.W. Albertson set the younger rangeman straight, as he now does his students.

46. Textbook example of true prairie- True prairie existing as a consociation of little bluestem with tall or composite dropseed as the assocaite species. The major climax forb was silverleaf scurfpea (Psoralea agrophyllum) followed by the similar slimflower scurpea or wild alfalfa (P. tenuifolia) which was the most common forb. Several plants of wavyleaf thistle (Cirsium undulatum) were present (see below). Eurasian grass species (invaders) were limited to small local areas of disturbance such as along cow paths. This simple (in both structure and species composition) climax grassland community was representative of the once widespread true prairie as described by Clements (1920, ps.121-131; Weaver and Clements, 1938, ps. 518-520, and Dodd [in Gould and Shaw, 1983, ps.346-348]).

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

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

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

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

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

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

49. Ultimate expression of true prairie- Such a superb example of true prairie that two more photographs of this now-relatively rare range cover type just had to be presented to students who might otherwise never know what true prairie really is. On a rolling upland in the Souther High Plains a simple "two-way mix" of little bluestem and tall dropseed (dominant and associate, respectively) comprised most of the range plant community of climax true prairie. Silverleaf and smallflower scurpea, wild alfalfas, along with wavyleaf thistle added a forb component.

The range vegetation presented in these two photographs was in Excellent range condition class. It was being grazed by black baldy cows and their calves. The relatively rarity of what was once a huge band of transition grassland that was an ecotone between mixed prairie and tallgrass prairie (Clements, 1920, ps.120- 131; Weaver and Clements, 1938, ps. 518-520) made these images at the top of this author's list of "trophies". Truly magnificant grassland! And picture-perfect management of this range. The highest praise to this land-owner and range manager.

50. Treasure found and shared- Turf of true prairie comprised almost exclusively of little bluestem and tall dropseed. There was some standing dead plant material of last year's herbage, but most standing crop was current year's growth. This range was being grazied by typical black baldy cows (1000-1200 pounds) and their calves. Range vegetation in these two "photoplots" was in Excellent range condition class.

This was bunchgrass prairie. It was grassland composed of cespitose species the genotypes/ecotypes of which had mature heights as to be more midgrasses than tallgrasses. Weaver and Bruner (1954) reported decreasing heights of maturity of little bluestem in the transition between tallgrass through true to mixed prairies. True prairie is the ecotonal grassland located between the more mesic tallgrass prairie to the humid and subhumid east (as in the Flint Hills, Osage Questas, Cherokee Prairie) and mixed prairie to the semiarid west (primarily the Great Plains). Perhaps the single most obvious feature of true prairie across much of this association is dominance by little bluestem versus big bluestem and Indiangrass although these latter two are typically present though ofen confined to more mesic microsites (Weaver et al., 1940; Weaver, 1943; Weaver and Bruner, 1954). Even within the zone of tallgrass prairie there is the little bluestem type and little bluestem consociation (Weaver and Fitzpatrick, 1934). Regional (zonal) dominance by little bluestem is key to distinguishing true prairie from tallgrass prairie.

Distinguishing distinction: the range vegetation presented in this and the two preceding two-slide sets was true prairie, the climax range type that is transitional and intermediate between tallgrass prairie and mixed prairie as was just explained. This was not degraded tallgrass prairie; it was the potential natural vegetation (high good to Excellent range condition class). Big bluestem, Indiangrass, and switchgrass are not the potential dominant decreasers of this range type or of this range site. Rather for this range site of the true prairie type (association) little bluestem, the least mesic (= most xeric) of the Four Horsemen of the Prairies, is the potential climax dominant (or co-dominant) with tall dropseed as the potential climax associate (or co-dominant). Tall dropseed is a decreaser not an increaser on this range site of true prairie, and tall dropseed is a dominant of true prairie (Weaver and Bruner, 1954, ps. 120, 123). Range site descriptions in the Norton County, Kansas Soil Survey (Soil Conservation Service) and Web Soil Survey (Natural Resources Conservation Service) were dead wrong. This range plant community was little bluestem-tall dropseed true prairie not big bluestem-Inidangrass tallgrass prairie.

The ease with which the more open turf and sod of bunchgrass prairie could be plowed with earlier types of steel moldboard plows and steam engines during the Homestead Era doomed almost all of the true prairie. Today, true prairie is one of the most endangered native plant communities in North America. School children raise money to save a single tree. The Nature Conservancy, government agencies, and universities cooperate to classify, locate, and save countless natural plant and animal communities as best they can. Even with such dedicated and zealous action that has unquestionably saved many a priceless piece of our natural heritage, true prairie remains unappreciated and, in fact, largely unrecognized. This is in spite of the devoted pioneeering ecological studies by the great prairiemen like John Weaver, Frederic Clements, Frederick Albertson and their many students.

For example, under combined efforts of the Natural Heritage Program, The Nature Conservancy, universities, and various other partners there are several state-level (statewide) vegetation classifications (basically the Braun-Blauget phytosocioloical system although usually not directly stated). In both the vegetation classification of Kansas (Lauver et al., 1999) and of Oklahoma (Hoagland, 2000) there was a Schizachyrium scoparium-Bouteloua cuirtipendula herbaceous alliance, but there should have been--and was not-- a Schizachyrium scoparium-Sporobolus asper and/or Schizachyrium scoparium-Sporobolus heterolepis herbaceous alliance. Even more telling, perhaps, was the omission of Sporobolus heterolepis and S. asper from all lists of "other species" under the Kansas community types led by Schizachyrium scoparium (Lauver et al., 1999, ps. 434-436)! Kuchler (1974) in effect missed the true prairie. Subsequent classifications followed suit.

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

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

52. Sward or turf of true prairie- Physiogonomy and species composition of true prairie co-dominated by little bluestem and tall dropseed. These two grass species composed over nine-tenths of the apparent cover and biomass of this range plant community. Plants of both species were in pre-boot stage, but had achieved complete leaf growth. This range was being grazed by cows and calves under light stocking. There was only limited herbaceous material from the previous year. Range condition class presented in this "photoquadrant" was Excellent.

53. More variation with more grazing- Local patch of past heavy grazing on the true prairie range in the Southern High Plains. Some spot grazing had taken place in the past on this range, but that pattern of grazing was not current (on-going). The local range vegetation was evidence of patchy heavy grazing in the recent past. The low turf (grass shoots) was not from present grazing, but was actually ungrazed or only lightly grazed plants of buffalograss and blue grama. These two sod-forming, shortgrass species accounted for almost all of the low-growing foliage although there was one small plant of Kentucky bluegrass "mixed in". Past heavy grazing had allowed shortgrasses to compete with little bluestem and tall dropseed so as to produce this mat of sod within clumps of the tall bunchgrasses. There were two or three plants of cespitose little bluestem. Also present were two plants (one large; one much smaller) of scurfpea (Psoralea agrophyllum) and four plants of wavyleaf thistle (Cirsium undulatum). All plants but the diminutive Kentucky bluegrass were natives.

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

54. Wavin' at ya'- Two plants of wavyleaf thistle (Cirsium undulatum) on true prairie dominated by little bluestem with tall dropseed as the associate species in the Southern (High) Great Plains. This was on the relict true prairie presented and described immediately above. Both of these plants (first slide) were at peak standing crop with one (on left) nearing senescence.Both plants had some fully open and some still unopened heads. Head (the more scientific name is capitulum, plural, capitula) is the Compositae inflorescence type in which there are sessile flowers, florets to be precise, grouped on a common receptacle known as a disk that is surrounded by an invloucre of bracts known as phyllaries (Smith, 1976, ps. 214, 291). Phyllaries are spine-tipped as, for that matter, the lobed leaves.

The second of these two slides presented a third plant that was also at peak growth (size) and full-bloom.This third wavyleaf thistle was featured in the next two-slide set which featured the capitulatum of this species.

Wavyleaf thistle is a native forb. It is not usually a weed, but instead an incidental to occasional species with sparsely populated plants. Presence of wavyleaf thistle on this mint condition true prairie attested to that typical situation (typical for pristine, not overgrazed) grassland.

Norton County, Kansas. Mid-June; full-bloom phenological stage.

55. Wavin' on true prairie- Heads (both open and still-to-open) of wabyleaf thistle (Cirsium undulatum) on true prairie of little bluestem (the dominant) and tall dropseed, the associate) in Southern High Plains. This was the same plant as that introduced in the immediately preceding slide (second slide of the preceding set). The base of the head (capitulum) below the disk is vaselike and covered with shingle-shaped spines.

Flowers of wavyleaf thistle, like those of other thistle species (Cynareae tribe in general), are a favorite of various insect species.

Norton County, Kansas. Mid-June; full-bloom phenological stage.

Passing note: an interesting phenomenon was shown by Norton County, Kansas true prairie. The invading Eurasian grass on true prairie is Poa pratensis whereas the main alien invader on northern tallgrass prairie is Bromus inermis and, on tallgrass to the east and south, Festuca arundinacea. Kentucky bluegrass is an invader of tallgrass (and true) prairie everywhere, but it was replaced as the number one exotic grass invader by smooth brome (in the north) and tall fescue (in the south). Kentucky bluegrass was introduced in North American grasslands (at least central ones) earlier in European settlement. This modification of dominant invaders began roughly in 1970s or 1980s within a few decades following introduction of Bromus inermis and Festuca arundinacea. All three Eurasian species are outstanding agronomic forage crops, and horrible weeds on native prairies. Textbook lesson in Weed Science.

56. Island of true prairie- Within the Mixed Prairie Region (mixed prairie was zonal or regional climax) a consociation of little bluestem had developed on a mesic north slope of a draw (a "canyon" in local idiom) holding forth as a postclimax. Higher up on and just below top of the "canyon" soapweed yucca (Yucca glauca) added a woody component so that this range vegetation could be interpreted as a savanna form (from perspective of physiogonomy). Conversely, if soapweed was viewed as s forb this vegetation would be strictly grassland of bunchgrass form comprising a strictly herbaceous form of true prairie.

In this population of little bluestem there were scattered plants of tall dropseed as the associate grass. There were very few plants of cheatgrass though enough to indicated the importing hand of the white man. The most common forb was western ragweed with scurfpea or wild alfalfa (Psoralea tenuifolia and/or P. agrophylla) as the second most abundant forb. This range had not been grazed by livestock for several years and resident wildlife had not removed much of the range herbage. There was considerable accumulation of standing dead plant material (necromass) along with current year's herbage (biomass). Maximum height of little bluestem shoots (again, ungrazed) was approximately two and a half feet so that this example of true prairie was more of a midgrass than tallgrass prairie. This intermediate height of little bluestem, which is usually a tallgrass species, was reflective of the transitional nature (from tallgrass to mixed prairies) of this grassland. Intermediate height (between midgrass and tallgrass species) and cespitose (tufted) habit (growth form) of dominants produced a bunchgrass physiogonomy that is charaacteristic of true prairie.

This range vegetation was a simple grassland plant community with no need for further elaboration. This relict tract spoke volumes by its mere presentation.

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

57. Another outstanding exmple of true prairie- Views of another true prairie range community (and another textbook example at that) in the High Plains. Tall dropseed and little bluestem were co-dominants with one of the other being locally more dominant. Indiangrass and sideoats grama were the associates. Indiangrass, the least mesophytic of the "big four" tallgrass species, constituted the element of this more eastward (and mesic) grassland and sideoats grama, probably the most widespread of the midgrasses, represented mixed prairie, the regional climax, to the west. Wavyleaf thistle was the major forb, but it was present at only trace cover. Plains pocket gopher (Geomys bursarius) were common on this range resulting in conspicuous low mounds on the land surface (eg. middle slide in this set).

This relict tract of approximately 20 to 30 acres was on a steep upper slope that even the most optimistic farmer realized could not be"broke out" without land loss.This climax vegetation was not being grazed by livestock at time of photographs (and obviously had not been for the last few yearts) or under drought conditions so that the range plant community exhibited structure, species composition, and maximum biomass of this cover type under minimum stress. Made for a beautiful picture of textbook true prairie.

Thomas County, Kansas. July (estival aspect). FRES No. 39 (Prairie Ecosystem). True prairie was not described by Kuchler, SRM, or Brown et al, 1998, all of whom should read the above remedial lesson (or read the classic literatue written by grassland ecologists who described climax vegeattion when there was more virgin sod to study). Western High Plains- Flat to Rolling Cropland Ecoregion 25d (Chapman et al., 2001).

58. Sward and composition of relict true prairie- Tall dropseed and little bluestem were co-dominants on a relict stand of true prairie in the Central High Plains with Indiangrass and sideoats grama, the associates, as third and fourth most abundant species. In the first of these two photographs tall dropseed had slightly more cover and general abundance than little bluestem. Two plants of Indiangrass and at least eight inflorescences (each loaded with ripe spikelets) of sideoats grama were conspicuous in the second photograph.

Sward refers to turf or the herbaceous layer(s) of vegetation in a plant community, especially of grasslands and savannahs. The sward of true prairie is generally comprised primarily of bunchgrass (cespitose or tufted) species more than sod-forming species. This bunchgrass feature was a major reason why the virgin sod of true prairie was typically broken with relatively greater ease than grasslands having greater proportions of sod-forming grasses. Hence, grasslands of the true prairie region were some of the first to be converted to cropland, and the grassland association most completely destroyed early on in the settlement phase (Clements, 1920, 121-122).

Once again students- The range vegetation presented in the last two sets of slides and captions was true prairie, a relict sample of the climax grassland association, the regional climax, that developed to the east of mixed prairie and west of tallgrass prairie (Clements, 1920, ps.121-133; Weaver and Clements, 1938, ps.518-521). This relict range plant community was not a degraded tallgrass prairie that "should have been" Indiangrass and big bluestem. It was textbook true prairie as described in the second sentence devoted to this Clementsian association in Weaver and Clements, 1938, p. 518). Little bluestem and tall dropseed were climax dominants while Indiangrass and sideoats grama were associate species that are dominants in tallgrass prairie and mixed prairie, respectively.

It was remarked here (as above) that true prairie was not recognized by any of the recent vegetation classifications. True prairie (as shown here) is real. It still exist in relict tracts like those presented. Failure to recognize this ecological fact is a deficiency or shortcoming of contemporary classifictions of potential natural vegetation.

A Taste of True Prairie in the Western Tallgrass Prairie Region

True prairie was first recognized by Clements (1920, ps. 121-131) and later established as textbook knowledge in the classic Plant Ecology (see esp. the second edition: Weaver and Clements, 1938, ps.518-520) as a transition grassland, "a broad ecotone" that constitutes "a fairly distinct belt between the tall-grass and mixed prairies..." (Weaver and Clements (1938, p. 519). This transition grassland was defined by Clements (1920, p. 122) as the Stipa-Koeleria Association that is composed of ranker-growing grass species from Andorpogon-dominated tallgrass prairie to the more mesic east and midgrasses from the Stipa-dominated mixed prairie to the less mesic west.

The range plant community featured in this short section did not have any Sporobolus species (at least not as dominants, associates, or even indicators) in which respect this range vegetation did not include the complete compliment of genera as specified in later refined descriptions of this association (Weaver and Clements, 1938, ps. 518-520). However, little bluestem, needle-and-thread, and Junegrass as the dominant, associate, and next major species, respectively, along with very limited cover of blue grama, the major shortgrass species, and almost no big bluestem and prairie sandreed, the two tallgrass species that are defining dominants of the Nebraska Sandhills, unequivocally placed this as transition grassland that was a "true" composite of an adjoining major range type (form of prairie) on either adjoining longitatudinal boundary. Substantial representation of three of the five consociations (Andropogon scoparius, Stipa comata, and Koeeleria cristata) in the true prairie, the Clementsian association, (Clements 1920, ps. 121-122) as the major three species in this range vegetation clearly defined this as an "island" of true prairie in a "sea" of the postclimax tallgrass (bluestem-prairie sandreed) prairie that comprises almost all of the Nebraska Sandhills other than the most westward, and less mesic portions. Also, presence of Junegrass as the least important of the three major species was consistent with the description by Clements (1920, p.122) that Junegrass was generally the least abundant (though the most widespread) of the dominants of true prairie.

From the perspective of potential natural vegetation units of Kuchler (1964, 1966) and Society for Range Management (SRM) rangeland cover types (Shiflet, 1994), which were based on Kuchler units, the true prairie range shown here was also ecotonal vegetation being a transition between Kuchler unit 67 (Nebraska Sandhills Prairie), which was SRM 602 (Bluestem-Prairie Sandreed), and Kuchler unit 60 (Wheatgrass-Bluestem-Needlegrass), which was SRM 606 (Wheatgrass-Bluestem-Needlegrass). Both Kuchler (1964, 1966) and the Society for Range management (Shiflet, 1994) failed to map and describe, respectively, true prairie. Omission of true prairie--one of the most clearly defined, verified, and historical (traditional) units of climax grassland in North America--by Kuchler (1964, 1966) has been a decades'-old mystery to the author of Range Types. It was completely understandable that authors of rangeland cover types for the Northern Great Plains section in Shiflet (1994, ps. 69-84) omitted true prairie (Stipa-Sporobolus-Andropogon-Koeleria) because they consistently followed Kuchler (1964, 1966)--even used his common name (English) titles--for all natural vegetation in that region. How Kuchler missed such a major unit of climax (potential natural) vegetation continued to baffle the current author.

In the present web publication this chapter, True Prairie, was included to recognize this major and distinctive North American grassland even though it had been mostly destroyed even by the time it was first recognized and described by Clements (1920, p. 121). And, no (in anticipation of the obvious question) that is not the reason why Kuchler (1964, 1966) omitted a unit entitled True Prairie, Needlegrass-Dropseed-Bluestem-Junegrass (Stipa-Sporobolus-Andropogon-Koeleria). He recognized the "extinct" California Steppe (Stipa) and that unit of potential natural vegetation was displaced permanently (in human time scale) by the naturalized unit, California annual grassland. Currently there are far more acres of Stipa-dominated true prairie than of Stipa-dominated California or Pacific bunchgrass prairie. No, Kuchler (1964, 1966) just missed it, plain and simple; and nobody else appears to have caught the error (or had the moxey to point out the mistake).

The example of true prairie treated in this short section showed the continuum-like arrangement and nature of grasslands in central North America, specifically that within the Nebraska Sandhills there is a transition (an ecotone) between postclimax tallgrass and mixed prairie. The rather limited mixed prairie in the western Nebraska Sandhills lacks a meaningful tallgrass component with these species occurring only as isolated individuals. For that reason, Sandhills mixed prairie was shown only in the Mixed Prairie chapter with no examples included under Tallgrass Prairie.

This was an isolated example of true prairie from the Nebraska Sandhills. It was clearly range vegetation open to interpretation..As was explained above that true prairie was first recognized by Clements (1920, ps. 121-131) and later included in the classic Plant Ecology (Weaver and Clements, 1938, ps.518-520) as a transition grassland, "a broad ecotone" that constitutes "a fairly distinct belt between the tall-grass and mixed prairies..." (Weaver and Clements (1938, p. 519). This transition grassland was defined by Clements (1920, p. 122) as the Stipa-Koeleria Association that is composed of ranker-growing grass species from Andorpogon-dominated tallgrass prairie to the more mesic east and midgrasses from the Stipa-dominated mixed prairie to the less mesic west.

This example of that climax ecotonal grassland from the Nebraska Sandhills was shown in the next two photographs. The range plant community featured in those up-coming slides did not have any Sporobolus species (at least not as dominants, associates, or even indicators) in which respect this range vegetation did not include the complete compliment of genera as specified in later refined descriptions of this association (Weaver and Clements, 1938, ps. 518-520). However, little bluestem, needle-and-thread, and Junegrass as the dominant, associate, and next major species, respectively, along with very limited cover of blue grama, the major shortgrass species, and almost no big bluestem and prairie sandreed, the two tallgrass species that are defining dominants of the Nebraska Sandhills, unequivocally placed this as transition grassland that was a "true" composite of an adjoining major range type (form of prairie) on either adjoining longitatudinal boundary. Substantial representation of three of the five consociations (Andropogon scoparius, Stipa comata, and Koeeleria cristata) in the true prairie, the Clementsian association, (Clements 1920, ps. 121-122) as the major three species in this range vegetation clearly defined this as an "island" of true prairie in a "sea" of the postclimax tallgrass (bluestem-prairie sandreed) prairie that comprises almost all of the Nebraska Sandhills other than the most westward, and less mesic portions. Also, presence of Junegrass as the least important of the three major species was consistent with the description by Clements (1920, p.122) that Junegrass was generally the least abundant (though the most widespread) of the dominants of true prairie.

In the present web publication a separate chapter, True Prairie, (under the Grassland biome) was included to recognize this major and distinctive North American grassland even though it had been mostly destroyed even by the time it was first recognized and described by Clements (1920, p. 121). And, no (in anticipation of the obvious question) that is not the reason why Kuchler (1964, 1966) omitted a unit entitled True Prairie, Needlegrass-Dropseed-Bluestem-Junegrass (Stipa-Sporobolus-Andropogon-Koeleria). He recognized the "extinct" California Steppe (Stipa) and that unit of potential natural vegetation was displaced permanently (in human time scale) by the naturalized unit, California annual grassland. Currently there are far more acres of Stipa-dominated true prairie than of Stipa-dominated California or Pacific bunchgrass prairie. No, Kuchler (1964, 1966) just missed it, plain and simple; and nobody else appears to have caught the error (or had the moxey to point out the mistake).

This example of true prairie showed the continuum-like arrangement and nature of grasslands in central North America, specifically that within the Nebraska Sandhills there is a transition (an ecotone) between postclimax tallgrass and mixed prairie. The rather limited mixed prairie in the western Nebraska Sandhills lacks a meaningful tallgrass component with these species occurring only as isolated individuals. For that reason, Sandhills mixed prairie was shown only in the Mixed Prairie chapter with no examples included under True Prairie.

59. Range vegetation open to interpretation- Nebraska Sandhills climax range vegetation that was interpreted as true prairie. This range plant community was dominated by little bluestem with needle-and-thread as associate species. Junegrass was a "distant third" major species followed even further by blue grama. There was trace cover of naturalized Japanese chess or Japanese brome. The only forb of consequence in this seasonal society was the native perennial composite, scaley blazing start or scaley gayfeather (Liatris glabrata). This combination of tall-, mid-, and even some shortgrass species was interpreted as true prairie, an ecotone between adjacent bluestem-prairie sandreed tallgrass prairie to the east and adjacent needla-and-thread--blue grama--western wheatgrass mixed prairie to the west. The range vegetation presented here was immediately west of the Nebraska Sandhills postclimax tallgrass prairie and also immediately east of Sandhills mixed prairie, this latter of which is the westernmost and driest of the various grasslands of that landform-soil complex.

The range plant community of this climax (Excellent range condition class) ectonal grassland was simple in species composition, including only four major native perennial grasses, yet relative diverse in architecture consisting of at least four layers of vegetation (counting the ground, soil surface, layer).

The introduction immediately before these two photographs described the historical interpretation of true prairie and explained why the currrent author included this short treatment Nebraska Sandhills true prairie in this Tallgrass Prairie (Interior) chapter. The combination of tallgrass and midgrass species and predominately cespitose growth form with substantial cover of sod-forming grass (mostly blue grama and some rhizomatous little bluestem) with a predominant bunchgrass prairie physiogonomy also was consistent with an interpretation and description of true prairie. In current (contemporary) view whereby true prairie is ignored or omitted this climax grassland could be described as the least mesic (most xeric) form or expression of bluestem-dominated tallgrass prairie. Again, it was explained above why the current author opted for the true prairie interpretation while at the same time having seen benefit to including this range vegetation here as an "vegetational bridge" between tallgrass and mixed prairies.

This range was upheld as the ultimate in wise use conservation of range resources. It was used as a perfect example of proper grazing management of rangeland, of stewardship beyond criticism. This photographer has seen his share of pristine vegetation, and none anywhere was any better than the "picture-perfect" model presented here. Furthermore, these two photographs were taken in early summer long before little bluestem would be at peak standing crop, and at end of a five- to six-year drought. Portrayal of this range at this aspect did show needle-and-thread at peak herbage crop to illustrate both the warm-season dominant (little bluestem) and the cool-season associate (needle-and-thread), along with the defining cool-seson Junegrass to good advantage.

Note standing dead herbage (residue or mulch) from the previous growing season. This is a sure sign of conservative stocking with cows and calves. It is essential for survival during severe (and worse) drough ts to stock on the light side because it is not known when the drought will break. Standing herbage, even if dead and weathered, is a savings account on the range. It amounts to that much hay and other fodder that this ranchman will not havee to purchase at times when such feed will, inevitably, be high-priced.

A well-deserved tip of the hat to this privte landowner and/or the range manager.

Cherry County, Nebraska. Late June; early estival aspect. FRES No. 38 (Plains Grassland Ecosystem) or No. 39 (Prairie Ecosystem), or a transition or combination of these. No Kuchler units or SRM rangeland cover type. This range type was in the general region of K-67 (Nebraska Sandhills Prairie) and SRM 602 (Bluestem-Prairie Sandreed), but it is transitional to K-60 (Wheatgrass-Bluestem-Needlegrass) and SRM 606 (Wheatgrass-Bluestem-Needlegrass). No unit in Brown (1998) either. Nebraska Sand Hills- Sand Hills Ecoregion, 44a (Chapman et al., 2001).

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

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

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

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

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

61. Little bluestem reigned solely supreme- At the margin of the Southern High Plains and Smoky Hills just a few miles distant from the tall dropseed -dominated true prairie (described immediately above) a consociation of little bluestem had developed on a shallow upland environment underlaid by caliche. There were few other grass species present though the author did encounter some sideoats grama and Japanese chess (Bromus japonicus). Of far greater abundance (and prominence) were numerous species of forbs including wild alfalfa or slimflower scurpea, leadplant (Amorpha canescens), wavyleaf thistle, western ragweed (Ambrosia psilostachya), black sampson or coneflower (Echinacea angustifolia), purple poppymallow (Callirhoe involucrata), and green milkweed (Asclepias viridiflora). Some plants of the shrub, western snowberry (Symphoricarpos occidentalis) were present.

Part of the Tertiary Ogallala formation remained on this rangeland as was seen prominently in the second slide.

Phillips County, Kansas. Late June-early estival aspect. Given published classifications this range vegetation would have the following designations (which did not accurately indicate the true prairie community): FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM 601 (Bluestem Prairie), generally, or, probably more specifically, SRM 710 (Bluestem Prairie). Bluestem "Tall-Grass" Series, 142.11, of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Central Great Plains- Rolling Plains and Breaks Ecoregion 27b (Chapman et al., 2001).

Note on equipment: these two photographs were taken within a few minutes of each other of the same true prairie range vegetation and under an identical sky ( light conditions). The slides (Provia 100F) were (are) identical in coloration. The glaring differences is color seen here were due 100% to the lack of precision in scanning by the very imprecise equipment, Epson Perfection 700. These two slides were placed one after the other (the second slide as shown here immediately after the first slide)in the Epson scanner. The scanner FUBARED the first slide and did the second one exactly right. Improper automatic scanning by this inferior equipment appeared to be at random, but for unknown reasons a rescanning of such slides typically resulted in the same--or worse--scanning. Warning to would -be purchasers: do not buy Epson equipment.

52. Some "forbaceous" characters profiled- Sward of the little bluestem-dominated true prairie introduced in the two preceding slides. These two progressively closer-in "photplots" featured some of the numerous forb species growing in this shallow upland range plant community. Leadplant, green milkweed, wild alfalfa or slimflower scurfpea, and western ragweed were visible in the first photograph while green milkweed and wild alfalfa were profiled in the second photograph. There was some sideoats grama as readily determined in the seond photogrpah by conspicuous "wood-shaving" leaves.

63. Winecups on true prairie- Purple poppymallow or winecups (Callirhoe involucrata) on the little bluestem-dominated true prairie featured immediately above. C. involucrata has a species range extending from Florida westward to Mexico, nnorth to the plains of North Dakota and westward to Arizona and southwestern Oregon.

Phillips County, Kansas. Late June, and obviously in full-bloom phenology.

True Prairie in the Northern Great Plains

True prairie probably remains at its greatest acreage and largest comparative proportions (relative to existing range vegetation) in the Northern Great Plains Region (Northern and upper Southern Great Plains and along margins of adjacent physiographic provinces). Clements (1920, p. 122) described true prairie as being at its greatest longitudinal width, seven degrees, at the forty third parallel of latitude and tapering to a longitude of one or two degrees in its northern and southern extremities. There were vegetational "islands" of true prairie to the west of the main body of true prairie, especially in the transition zone (ecotone) between the Palouse Prairie of the Pacific Northwest and Great Plains grasslands (Weaver and Albertson, 1956, ps. 339-341). (This was described in greater detail below.) Remnants--sometimes quite large ones--of this transitional grassland are to be found as isolated grasslands within the overall Northern Mixed Prairie.

Subsequent and more detailed studies, as summarized by Barker and Whitman (1989) and incorporated into rangeland cover types (Shiflet, 1994), indicated that much of the climax grassland in the Northern Great Plains lacks both a dominant shortgrass component and a functional tallgrass component (when little bluestem exist as a midgrass). These grassland dominant (cover types) fit the classic description/designation of true prairie. Such grasslands include the following rangeland cover types published by the Society for Range Management (Shiflet, 1994): SRM 606 (Wheatgrass-Bluestem-Needlegrass), SRM 607 (Wheatgrass-Needlegrass), and SRM 610 (Wheatgrass). Weaver and Albertson (1956, p. 340) followed the pioneer work of Clements (1920) and indicated that it was occurrrence of shortgrass species of Bouteloua (mostly B. gracilis) as a dominant that distinguished mixed prairie from true prairie. Another shortgrass species that occurs as a dominant (or major associate species) in the Northern Great Plains is Sandberg's bluegrass (Poa secunda=P. sandbergii). Adherence to the historic distinction between true and mixed prairies would designate the two related rangeland cover types (Shiflet, 1994) of SRM 608 (Wheatgrass-Grama-Needlegrass) and SRM 609 (Wheatgrass-Grama) as mixed prairie: Hence, SRM 606, SRM 607, and SRM 610 were included in this True Prairie chapter wheras SRM 608 and SRM 609 were treated in the chapter, Mixed Prairie. SRM 611 (Blue Grama-Buffalograss) was, obviously, covered in the Shortgrass Prairie chapter.

Climax true prairie in the vast domain of the Northern Great Plains (and parts of adjoining physiographic provinces) includes among its dominants such midgrass species as western wheatgrass (Agropyron smithii), needle-and-thread (Stipa comata), porcupinegrass (S. spartea), Junegrass (Koleria cristata), and sideoats grama (Bouteloua curtipendula) as well as tallgrass species like little bluestem, which in this region has mature shoot heights no taller (and, sometimes, shorter) than those of midgrass species. Both bunchgrass and sod-forming grass species are present with some species like little bluestem assuming both habits (Weaver and Clements, 1938, p. 518).

In the southern portion of the Northern Great Plains there is an ecotone of true prairie and tallgrass prairie that extends northward from the Nebraska Sandhills whereas another ecotone ot true prairie and mixed prairie is to be found in western portions (Cleements, 1920, p.122; Weaver and Clements, 1938, ps. 519-520; Weaver and Albertson, 1954, p. 318).

The definitive work on the natural vegetation of the Northern Great Plains remains that of Barker and Whitman (1989). Descriptions of rangeland cover types of the Northern Great Plains Region published by the Society for Range Management (Shiflet, 1994) were taken directly from Barker and Whitman (1989), but some plant communities in the earlier document did not make it into the SRM (Shiflet, 1994) publication.

The following section was devoted to true prairie in the Northern Great Plains. This included range vegetation of the White River and Little Missouri River Badlands except for the Rocky Mountain juniper (Juniperus scoparium) forest cover type (SAF 220) which was included in the chapter, Juniper-Pinon Woodland, Woodlands and Forests.

Some of the major gasslands (grassland cover types) in this vegetational region of the Northern Great Plains were interpreted in this publication as true prairie because the range cover (dominance) types presented and described did not have a major shortgrass component. Kuchler (1966) showed the potential natural vegetation for this region as Grama-Needlegrass-Wheatgrass (K-57), but previously Kuchler (1964, ps. 64, 66) distinguished between vegetation units of Grama-Needlegrass-Wheatgrass (Bouteloua-Stipa-Agropyron) and Needlegrass-Wheatgrass (Stipa-Agropyron). This same distinction was followed by Barker and Whitman (1989) and in Shiflet (1994). Rosiere (herein) regarded the Grama-Needlegrass-Wheatgrass unit as mixed prairie and the Needlegrass-Wheatgrass unit as true prairie. This distinction (difference between these two designations) by the present author seemed to be most consistent with tradition or classical treatments beginning with Clements (1920, ps.121-131,135-137) and Weaver and Clements (1938, ps. 518-520, 523-525).

Kuchler (1964, p. 67) also recognized a Wheatgrass-Bluestem-Needlegrass (Agropyron-Andorpogon-Stipa) unit.of grassland. This unit seemed even more obviously to be true prairie. Again this was consistent with the original designation of true prairie (Clements, 1920, ps. 121-131) and of the distinction between true prairie and mixed prairie (Weaver and Clements, 1938, ps. 518-520, 523-525). It should be stressed that in this region (Northern Great Plains) 1) little bluestem is of a shortened height so that it is more of a midgrass than a tallgrass species and 2) a shortgrass dominant is lacking. Blue grama is present even in the Needlegrass-Wheatgrass (Stipa-Agropyron) unit, but blue grama, buffalograss, hairy grama, etc. are not even associate--let alone, dominant--species in these grasslands. In the original designation of true prairie and mixed prairie it was absence and presence, respectively, of blue grama and/or buffalograss as one of the dominants that served as basis of this distinction (Clements, 1920, ps. 121-122, 135-137).

Note on organization/location: Grasslands of the Wheatgrass-Grama-Needlegrass rangeland cover type (SRM 608; Shiflet, 1994) are obviosuly mixed prairie and thus were treated under the Mixed Prairie chapter under the Grasslands heading in Range Types of North America.

Further note on organization/location: woodlands of green ash (Fraxinus pennsylvanica) and Rocky Mountain juniper (Juniperus scoparium) in the Northern Great Plains were included in chapters Juniper-Pinon Woodland and Central and Southern Forests-I under Woodlands and Forests, respectively.

Note: In addition to true prairie in the Northern Great Plains there are immense acreages of grazing disclimax grassland comprised of 1) varying mixes of Kentucky bluegrass (Poa pratensis) and smooth brome (Bromus inermis) and, at the ultimate state of disturbance (primarily overgrazing) climax, 2) monocultures of Kentucky bluegrass. For example, Kronberg et al. (2011) described range on the Northern Great Plains Research Center near Mandan, North Dakota on which Kentucky bluegrass and smooth berome predominated over blue grama, green needlegrass, porcupinegrass, and western wheatgrass. The dominant forb was yellow sweet-clover (Melilotus officinalis).

These two disclimax range types made up of 1) various combinations of Kentucky bluegrass and smooth bromegrass and 2) single-species stands of Kentucky bluegrass are naturalized types much like the California annual grassland type and cheatgrass or downy brome (Bromus tectorum) type. The Kentucky bluegrass and Kentucky bluegrass-smooth brome types were not recognized as such by authors in Shiflet (1994). These two disclimax grassland types are comprised of exotic (Eurasian in origin), perennial grass species that were intentionally introduced as agronomic crops by the white man in contrast to Eurasian annual grasses that were inadvertently brought over by Christopher Columbus and his enlarged clan from Europe and the British Isles. Eurasian disclimax grasslands as well as skilfully managed pastures and hay fields of smooth bromegrass and Kentucky bluegrass were covered under the chapter, Introduced Forages, Grasslands.

64. Little Missouri River Badlands- Grand landscape-scale views of badlands of the Little Missouri River with a sweeping mosaic of Northern Great Plains grasslands of several rangeland cover types along with forest cover types including Rocky Mountain juniper (Juniperus scoparium) woodlands, quaking aspen (Populus tremuloides) groves, and floodplain forests of eastern cottonwood (Populus deltoides), willow (Salix spp.), and green ash (Fraxinus pennsylvanica).

True prairie dominated by needle-and-thread and western wheatgrass--SRM 607 (Wheatgrass-Needlegrass)--was the range vegetation featured in foreground of both of these slides. There were some plants of silver sagebrush (Artemisia cana) in this small patch of true prairie.

Theodore Roosevelt National Park (North Unit), McKenzie County, North Dakota. Late June, late vernal to early estival aspect and society (peak standing crop of cool-season grasses). Various range plant communities and range types. FRES No. 38 (Plains Grasslands Ecosystem). K-59 (Wheatgrass-Needlegrass). Mosaic of SRM types 606, 607, and 610. Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40). No FRES No. or K-unit, but there was SAF 239 (Pinyon-Juniper). FRES No. 17 (Elm--Ash-Cottonwood Forest Ecosystem). K-89 (Northern Floodplain Forest). SAF 235 (Cottonwood-Willow). FRES No. 19 (Aspen-Birch Forest and Woodland Ecosystem). K-97 (Northern Hardwoods, Seral Stages). SAF 217 (Aspen). Northwestern Great Plains- Little Missouri Badlands Ecoregion Ecoregion 43b (Byce et al., Undated).

65. Swatch of wheatgrass- Close-up of sward of western wheatgrass that developed on precipice of Little Missouri River Badlands. This was a "photoplot" of the wheatgrass-needlegrass true prairie introduced in the immediately preceding slide.

Theodore Roosevelt National Park (North Unit), McKenzie County, North Dakota. Late June, late vernal to early estival aspect and society (peak standing crop of cool-season grasses). Various range plant communities and range types. FRES No. 38 (Plains Grasslands Ecosystem). K-59 (Wheatgrass-Needlegrass). Mostly SRM 607 (Wheatgrass-Needlegrass). Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40). Other range types, complete with FRES, K- units, and SAF designations in distant (and mostly not discernable) far background were given in the immediately preceding caption. Northwestern Great Plains- Little Missouri Badlands Ecoregion Ecoregion 43b (Byce et al., Undated).

66. True prairie (and trees) above the Little Missouri- Landscape on the outer margin of the Little Missouri River Badlands. This climax range vegetation was a mosaic of 1) true prairie consisting of various plant communities including wheatgrass-bluestem-needlegrass, wheatgrass-grama-needlegrass, and wheatgrass-needlegrass cover types, 2) grass-shrub (mostly Artemisia cana) savanna, 3) groves and gallery forests of green ash and eastern cottonwood along drainages (eg. center midground of this photograph) and 4) woodlands of Rocky Mountain juniper (distant background of this slide in hillside draws).

The major range plant community in this photograph (foreground and majority of midground) was the wheatgrass-bluestem-needlegrass cover (dominance) type.

This landscape-scale scene presented the physiography (topographic relief) of the outermost portion of L<ittle Missouri Badlands. Closer views of low buttes or high hills set against the background sky wre shown at shorter camera distance in the immediately following two slides.

Theodore Roosevelt National Park (South Unit), Billings County, North Dakota. Late June, early estival aspect and societies. FRES No. 38 (Plains Grasslands Ecosystem). K-59 (Wheatgrass-Needlegrass). Major rangeland cover type was SRM 606 (Wheatgrass-Bluestem-Needlegrass). Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40). Northwestern Great Plains- Little Missouri Badlands Ecoregion 43b (Bryce et al., Undated).

67. On the border of the Badlands- True prairie that developed at the outer edge of the Little Missouri Badlands.Grassland vegetation as range cover (dominance ) types was wheatgrass-needlegrass (SRM 606) with the three dominants of western wheatgrass, needle-and-thread, and green needlegrass within (surrounded by) the overall cover type of wheatgrass-bluestem-needlegrass (SRM 607). Other important grasses (local associates) were Junegrass, blue grama, and sdiedots grama. There were almost no forbs to speak of. There were sparcely spaced plants of silver sagebrush (Artemisia cana), but these widely isolated individuals were not of sufficient cover, density, general abundance to make up a woody component (ie. there was no savanna vegetation visible in these two photographs).

Physigonomy, structure, and a general view of species composition of this grassland vegetation was emphasized in thesee two photographs.

High hills or low buttes in background of the preceding sky were similar to the three (at least parts of three) shown in these two slides.

Theodore Roosevelt National Park (South Unit), Billings County, North Dakota. Late June, early estival aspect and societies. FRES No. 38 (Plains Grasslands Ecosystem). K-59 (Wheatgrass-Needlegrass). SRM 606 (Wheatgrass-Bluestem-Needlegrass). Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40). Northwestern Great Plains- Little Missouri Badlands Ecoregion 43b (Bryce et al., Undated).

68. Parts and patches- A wide-angle view in the first of these two photograph presented the combination of 1) landscape-scale and 2) range plant community structure and composition of a wheatgrass-bluestem-needlegrass true prairie in the Little Missouri River Badlands. Little bluestem grew in and dominated local areas (patches of tan- or light brown-colored straw that were remnants of last year's shoots) thereby forming little bluestem consociations. These consociations were scattered within other consociations and natural "mixtures" of such dominant and associate cool-season species as needle-and-thread, green needlegrass, western wheatgrass, and Junegrass. There was some cover of blue grama, but sideoats grama (second slide) had substantially greater cover and was the more important Bouteloua species. There was insufficient cover and density of blue grama to compose a shortgrass component. The abundant broad-leafed plants in foreground were prairie rose (Rosa arkansana).

The second slide was a closer-up view of range vegetation seen in the first slide so that the second was a "photoplot" that was "nested" within the first (larger-scale) photograph. The grassland vegetation seen in the second slide was that at edges of 1) a local community co-dominated by western wheatgrass and Junegrass with a nice plant of sideoats grama thriving with these festucoid grasses (foreground) and 2) a consociation of little bluestem (tan-colored straw behind in the midground). This true prairie made up of both cool-season (western wheatgrass, Junegrass) and warm-season species (little bluestem and sideoats grama) and of festucoid, eragrostoid (sideoats grama), and panicoid (little bluestem). The two major forbs in this second slide were lambstongue ragwort (Senecio intergerrimus) which was blooming and purple coneflower (Echinacea angustifolia) whch was in prebloom stage (not even elongating the sexual shoot).

In the Northern Great Plains Region little bluestem grows to a height and general size of a midgrass rather than that of a tallgrass species (the size it reaches in the Southern and much of the Central Plains). In the Northern Plains little bluestem grows to about the same height as sideoats grama. (This was why the second slide in this set featured a cespitose plant of sideoats grama. Admitttedly, this robust specimen of sideoats grama was in the foreground so that it appeared relatively tlaller, but it was growing beside Junegrass plants so viewers got get an accurate--and not a misleading--presentation or guage as to its height and general size. Two photographs were presented two slide-caption sets below in which little bluestem and sideoats grama were growing side-by-side--and to the same height.)

Clements (1920, p.122 ) listed little bluestem, western wheatgrass, and Junegrass as three of the five plant species forming consociations in the true prairie.

Theodore Roosevelt National Park (South Unit), Billings County, North Dakota. Late June, early estival aspect and societies. FRES No. 38 (Plains Grasslands Ecosystem). K-59 (Wheatgrass-Needlegrass). SRM 606 (Wheatgrass-Bluestem-Needlegrass). Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40). Northwestern Great Plains- Little Missouri Badlands Ecoregion 43b (Bryce et al., Undated).

69. A pocket near the top- A small sloughed bit of land (microsite) at the top of a lower hill on the outskirts of the Little Missouri River Badlands was home to creeping juniper (Juniperus horizontalis) amid a local consociation of litttle bluestem with sideoats grama as the associate species. Strictly speaking this was a shrub-midgrass savanna even though there was not a readily noticed physiogonomy characteristic of a grassland with sporadic cover of woody species. Recall from the immediately preceding caption that in this portion of the Northern Great Plains little bluestem assumes a height of midgrass and not a tallgrass species.

There were a couple of plants of lambstongue ragwort which was about the only forb in this range plant community.

Theodore Roosevelt National Park (South Unit), Billings County, North Dakota. Late June, early estival aspect and societies. FRES No. 38 (Plains Grasslands Ecosystem). K-59 (Wheatgrass-Needlegrass). SRM 606 (Wheatgrass-Bluestem-Needlegrass). Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40). Northwestern Great Plains- Little Missouri Badlands Ecoregion 43b (Bryce et al., Undated).

70. At its highest- True prairie dominated by little bluestem (sideoats grama was associate species) on the outer margin of Little Missouri River Badlands. This was a more mesic habitat on a northeast slope atop a lower hill among a series of such hills. Barker and Whitman (1989) labeled this or similar land shapes as "low knolls". Climax range vegetation presented in these two photographs was an unusually good example of the widespread little bluestem-dominated community that was part of the mosaic of the overall or general wheatgrass-bluestem-needlegrass range cover (dominance) type. (Examples of this general range dominance type were presented above.) Distinctive patches dominated by little bluestem with their characteristic tan- or brownish-colored shoots or "straw" are either local consociations of almost "pure" (single-species stands of) little bluestem or simple mixtures of little bluestem with relatively few other plant species. Sideoats grama is one of the more (often, the most) common of these other species. In fact, sideoats grama is generally the most common associate species affilitated with little bluestem on this range cover type.

Interestingly, and for whatever reason(s), these warm-season species--panicoid little bluestem; eragroastoid sideoats grama--usually grow together forming communities that are distinctively and conspicuously different from surrounding communities comprised of western wheatgrass, needle-and-thread, and green needlegrass. In the view of Landscape Ecology this spatial arrangement, structure, and physiogonomy can be regarded as little bluestem patches within a wheatgrass needlegrass matrix, the whole making up the overall wheatgrass-bluestem-needlegrass range plant community.

These two "photoplots" taken at farther and shorter (in that order) cameral distances showed that mature shoots of little bluestem and sideoats grama were generally of the same height. This is the result of little bluestem shoots only reaching a mature (flowering/fruiting) stature characteristic of midgrass species such as sideoats grama, western wheatgrass, and needle-and-thread. In fact, and as shown in both of these photographs, sideoats grama often grows to a height exceeding that of little bluestem (these slides provided side-by-side comparisons). This similarity in height is in contrast to habits found farther south and east, especially in tallgrass prairie such as in the Nebraska Sandhills, where little bluestem is a tallgrass species towering above sideoats grama. In other words, this similarity in height and general size of plants of these species is due to little bluestem growing shorter and smaller and not sideoats grama or western wheatgrass reaching greater heights.

Smaller--especially, shorter--size of little bluestem was one of the differences between true and tallgrass prairies. Little bluestem was the only species listed by Clements (1920, ps. 122, 132) as forming consociations in both true prairie and tallgrass prairie (ie. Andropgogn scoparius was the major species that proved the affinity of these two major grasslands, two cover or dominance range types). The shortness and generally smaller size of little bluestem (ie. its occurrence as a midgrass) was a key criterion used by the current author to designate the wheatgrass-bluestem-needlegrass rangeland cover type as true rather than as mixed prairie.

The major forb in this simple plant community was lambstongue ragwort. Plants of this composite species were shown at larger size (closer camera distance) in the first photograph. A shrub commonly associated with little bluestem-dominated vegetation in the Little Missouri River Badlands and adjoining areas was creeping juniper. (This phenomenon was shown in the immediately preceding, and several other, slides in this section.)

Theodore Roosevelt National Park (South Unit), Billings County, North Dakota. Late June, early estival aspect and societies. FRES No. 38 (Plains Grasslands Ecosystem). K-59 (Wheatgrass-Needlegrass). SRM 606 (Wheatgrass-Bluestem-Needlegrass). Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40). Northwestern Great Plains- Little Missouri Badlands Ecoregion 43b (Bryce et al., Undated).

71. Short savanna just short of the badlands- A rangeland community of the little bluestem-creeping juniper habitat type or creeping juniper/little bluestem dwarf shrubland (Hansen et al., 1984; Montana Natural Heritage Program, 2002) that developed on a ridge top at margin of Little Missouri River Badlands. This creeping juniper-prairie grass savanna was a midgrass variant in contrast to the mixed grass variant which was included in the chapter, Mixed Prairie I. Midgrass status was the situation because throughout the semiarid Northern Great Plains little bluestem attains height and relative size of a midgrass rather than that of a tallgrass as it is farther to the south and east. In the Northern Great Plains Region little bluestem is more the stature and size of sideoats grama and western wheatgrass. Clements (1920, p.122) recognized a little bluestem consociation for true prairie. This is that vegetational unit (consociation) with a co-dominant coniferous shrub so that this climax range vegetation was a midgrass-shrub savanna or, with a slightly different emphasis, a true prairie ecotone.

Three other grass species that, at most, were associates to little bluestem were plains reedgrass (Calamagrostis montanensis), green needlegrass, and plains muhly (Muhlembergia cuspidata). Plains reedgrass was the most abundant of these three species. Other important grasses included western wheatgrass and green needlegrass. There were numerous forbs, mostly composites. The most abundant of these was (Solidago rigida). Stiff goldenrod and little bluestem were co-dominants in certain locations such as that shown in foreground of the second of these slides.

Although there was enough cover of creeping juniper that this range vegetation could be interpreted as a savanna it could as easily be regarded as grassland (true prairie) with a sporadic woody component. Savannah status was largely drawn from adjacent range plant communities such as that on a neighboring ridge top (presented shortly below) as well as that found in the same area on which there was greater cover of creeping juniper (eg. examples presented above in this section). True prairie dominated primarily by little bluestem with isolated creeping juniper (or little bluestem-creeping juniper savanna) in the interior (on ridge tops) of the Little Missouri Badlands was more mesic than that shown from slopes or hill sides in this general area such as examples in the Golden Valley on which there were other associate species such as bluebunch wheatgrass, Junegrass, blue grama, and Sandberg's bluegrass. That range vegetation was mixed prairie-creeping juniper savanna. Though similar, those range plant communities were obviously mixed prairie as they had a substantial shortgrass component. Hence, that similar climax range vegetation was included in the Grassland chapter entitled Mixed Prairie-I.

In the background of the landscape shown here down in coulees and draws there were woodlands of two range types: 1) groves of green ash as the only tree species.and 2) tree stands dominated by Rocky Mountain juniper sometimes with green ash (Fraxinus pennsylvanica) as a minor species.

Note on grassland location: Given that in the strict sense those savannahs that were a mixed grass (as in mixed prairie)-savanna community or, same thing, mixed prairie with creeping juniper versus true prairie with creeping juniper were treated in chapter, Mixed Prairie I, in order that the mixed prairie form of the creeping juniper savanna was kept separate from the true prairie form of the creeping juniper savann. By contrast, both true and mixed prairie forms were retained together in the chapter, Mixed Prairie I, for students desiring to view them together and compare them more easily. The creeping juniper-grassland (more precisely, a grass-creeping juniper savanna)--which is a climax plant community --that is true prairie belonged here under the heading of true prairie (True Prairie chapter).

Note on woodland location: Green ash groves were included in the chapter entitled Southern and Central Forest while Rocky Mountain juniper woodlands were included in the Juniper-Pinyon Woodland chapter both within Range Types of North America.

Theodore Roosevelt National Park (North Unit), McKinzie County, North Dakota. Late June, early estival aspect and society. All warm-season grasses were still in early phenological stages. FRES No. 38 (Plains Grasslands Ecosystem). K-59 (Wheatgrass-Needlegrass). Savanna variant of SRM 606 (Wheatgrass-Bluestem-Needlegrass). Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40). Northwestern Great Plains- Little Missouri Badlands Ecoregion 43b (Bryce et al., Undated).

72. Another perspecitve of the ridge-top, short savanna- A short walk across a ridge top on the margin of Little Missouri River Badlands furnished viewers a pair of nested (one inside the other) "photoquadrants" of the range plant community introduced in the preceding pair of "photoplots". This climax range vegetation was a savanna form of true prairie, specifically of the range type (or subtype) designated as the little bluestem-creeping juniper habitat type or creeping juniper/little bluestem dwarf shrubland (Hansen et al., 1984; Montana Natural Heritage Program, 2002). This is basically a true prairie grassland with a climax shrub layer, hence a midgrass-short shrub savanna.

The first slide provided a smaller landscape view (from a slightly different camera loction) of the same range vegetation presented in the immediately preceding pair of photographs. The second of the two slides showed the sward of this true prairie savannah so as to provide an idea of its structure and composition. In this "photoplot" stiff goldenrod was co-dominant with little bluestem while creeping juniper was an associate species. Other major--locally, associate--species included plains muhly and plains reedgrass. Other important--though generally less widespread--species were western wheatgrass and green needlegrass. Thus, this grassland vegetation was a variant of the Wheatgrass-Bluestem-Needlegrass range type described by Barker and Whitman (1989) and Shiflet (1994).

The first of these two slides along with the preceding two slides showed physiogonomy of this climax plant community. Woodland vegetation showed in these three slides, especially the first one of this slide-caption set, was of two types: : 1) stands of green ash as the only tree species properly described as groves and 2)woodlands dominated by Rocky Mountain juniper sometimes with green ash (Fraxinus pennsylvanica) as a minor species. These woodland range communities developed in draws or coulees that were more mesic than ridge tops or south and west slopes.

73. Turf and turd- Sward of a little bluestem-creeping juniper savanna (or grassland with a major woody component) with Juniperus horizontalis conspicuous. Other grass species present included green needlegrass, the local associate species (to little bluestem and creeping juniper) in the first of these two slides, plains reedgrass, plains muhly, and, in the second slide, western wheatgrass. Plains reedgrass was also a local associate species on some (drier) microsites of this same savanna range. Also on this savanna range, and included in the second "photoplot", was threadleaf caric sdege.

The buffalo chips in the first of these two slides showed that this range was being grazed by North American buffalo (Bison bison) which was a (probably, the) dominant native herbivore and reminded students that 1) range is defined by use of native vegetation as pasture for grazing/browsing animals, 2) range ecosystems are designated by having consumers and decomposers (as those that rot buffalo dung) as well as producers (such as those species shown here), 3) recycling of nutrients is an essential and defining ecosystem function, and 4) herbivory consist of several impacts on range plants and range plant communities including covering (as with dung deposition) as well as removal by eating plant material (biomass that becomes forage).

"Oh, buffalo gals won't ya come out tonight ..." (For those of ya'll ignorant of frontier culture, that folk song reference was to the nightly duty of girls and women gathering buffalo chips for cooking and warmth. We had real she-folk in those times!)

Theodore Roosevelt National Park (North Unit), McKinzie County, North Dakota. Late June, early estival aspect and society. FRES No. 38 (Plains Grasslands Ecosystem). K-59 (Wheatgrass-Needlegrass). Savanna variant of SRM 606 (Wheatgrass-Bluestem-Needlegrass). Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40). Northwestern Great Plains- Little Missouri Badlands Ecoregion 43b (Bryce et al., Undated).

74. Another short (and more like a) savanna on another ridge top- Another pair of nested "photoplots" of a little bluestem-creeping juniper true prairie savanna that developed on an east slope of a ridge on the perimeter of the canyons of Little Missouri River Badlands. Range vegetation on this ridge had a higher proportion of creeping juniper and a more equal foliar cover of major grass species which included (in addition to the dominant little bluestem): plains muhly, plains reedgrass, green needlegrass, and western wheatgrass. Threadleaf caric sedge was also present throughout. The principal forb was stiff goldenrod as was the case for the ridge top little bluestem-creeping juniper savanna preented immediately above. In range vegetation on this adjoining ridge stiff goldenrod was not nearly as abundant as in the other ridge vegetation in which it was locally co-dominant with little bluestem.

Woody vegetation in draws (backgrounds of both photographs) was a combination of Rocky Mountain juniper and green ash as both juniper woodland with small cover of green ash and green ash stands that developed into groves.

75. Flat conifer and friends- "Photoquadrant" of a portion of a single Rocky Mountain juniper accompanied by--"believe it or not"--shoots of little bluestem, green needlegrass, plains muhly, western wheatgrass, stiff goldenrod, and an unidentified Lupinus species (in that rough order of rank based on apparent cover).

The abundant crop of fleshy seeds (cones) was apparent in this wide-angle (28mm lens) view.

76. Flat foliage and fleshy seeds- Needles and the spined, globular seeds of creeping juniper (Juniperus horizontalis).

Theodore Roosevelt National Park (North Unit), McKinzie County, North Dakota. Late June,

77. Creeping its own- Yellow wild-buckwheat (Eriogonum flavum) growing in the minst of bough of creeping juniper and little bluestem with some plains reedgrass and plains muhly. The Eriogonum species comprise one of the most widespread species of forbs, shrubs, and subshrubs (suffrutescent plants) on the Western Range. McGregor et al. (1986, ps. 215-219) described 14 species of Eriogonum for their interpretation of the general Great Plains Region.

E. flavum is, according to MdGregon et al. (1986, p. 216-217) restricted to the Great Plains, but this was incorrect as yellow wild-buckwheat is also native to North America west of the Rocky Mountains occurring in the Palouse Prairie and Northern Rocky Mountains. For example it is widespread in the Idaho Panhandle.

Theodore Roosevelt National Park (North Unit), McKinzie County, North Dakota. Late June, peak-bloom phenological stage.

78. Diversty on a ridge top- Sward of a little bluestem-creeping juniper savanna on east slope of a ridge on outskirts of canyons of Little Missouri River Badlands. In addition to the two co-dominants other plant species included plains reedgrass, the main associate, western wheatgrass, plains muhly, prairie rose (Rosa arkansana), and the very conspicuous pasture puffball (Lycoperdon craniformis= Calvita craniformis).

These two "photoquadrants" were obviouysly nesteed with the second being a zoomed in view of the pasture puffball (mushroom) and several shoots of prairie rose. Both of these species have extremely broad biological (species) ranges across North America. Pasture puffball, one of the most common mushrooms on North American grasslands has a global distribution. This is perhaps not surprising when one understands that tiny spores of the fungii are carried aloft so as to circle the planet. Prairie rose has a species range extending from Mexico north to the Canadian Prairie Provinces. This suffrutescent species is also a common grassland species.

Reducers or decomposers comprise one of four biotic components (one group of organisms) of range ecosystems. The reducers include microorganisms (bacteria and protozoa), fungi, and saprophytic vascular plants (saprophytes). Decomposers, like consumers, are heterotrophs. In contrast to autotrophs or self-nourishing organisms like chlorophyll-containing plants (producers), heterophytic organisms cannot produce food by photosynthesis or chemosynthesis. Heterotrophs depend on autotrophs (producers) for their food. The suffix "troph" refers to trophic levels, the "links" (steps) in food chains, the sequence by which energy in the form of food (feed) is transfered from producers to primary consumers through to the final (say, tertiary) consumers. Dead organic material of produders and consumers becomes the food of reducers (or decomposers, which get their ecosystem functional name from decomposition or the process of rotting). This dead organic material that is available for rotting is called detritis the plant portion of which is refered to as litter. "Troph" is derived from the Greek, trophe meaning nourishment. Saprophytes (saprophytic plants)--note the suffix, phyte meaning plant--get their nourishment from the organic matter of dead organisms (plant or animal). Saprophytes (saprotrophs is the all-encomposing term applied to all saprophytic organisms) include the fungi (kingdom, Fungi; plants in the two-kingdom system).

The fungi are often known by the layman's generic terms of "mushrooms" or "toadstools". These organisms (range plants in the lexicon of this rangeman) are some of the largest, most conspicuous, and curiosity pricking saprophytes in range ecosystems. Two examples of range or, in this case and more specifically, grassland (true prairie) "mushrooms" from little bluestem-creeping juniper savanna (the creeping juniper-little bluestem habitat type) where presented below for students of range ecosystems. Both of these species are of the class Hymenomycetes, subdivision of Basidiomycotina (Basidomycetes). (Pay attention: quiz at end of class.)

79. Ripening and ripe- Two carpophores or fruiting bodies of skull puffball (Lycoperdon craniformis= Calvatia craniformis) on little bluestem-creeping juniper savanna (a grassier form of creeping juniper dwarf shrubland) in Little Missouri River Badlands. Fruiting body at left was immature though ripening while a ripe or fully mature fruiting body was at right. Both carpophores were still attached to the land surface and shown in a dorsal or top-down view on the range. The carpophore or fruiting body (the sexually reproductive, spore-producing, organ or structure of a fungus) is the conspicuous or readily seen part of a fungus. The fruiting body t is what most people see (and therefore assume) as the "mushroom".

The next two slides-caption sets showed and briefly described these two carpophores.

The green plant in immediate left-center and in front of carpophores was prairie rose. Most of the grass shoots were of plains reedgrass and, secondly, western wheatgrass.

Theodore Roosevelt National Park (North Unit), McKinzie County, North Dakota. Late June.

80. Getting there- Top-down view of the immature fruiting bodies(carpophores) of skullcap puffball introduced in the preceding photograph. (This photograph was taken from the opposite direction of previous slide.) Unfortunately, this view did not present enough of a lateral (side) view to show the rounded skull-resenbling fruiting body of this species. Viewers will have to use their range imagination for this. Lycoperdon (Calvatia) species are in the Lycopodales or Hymenogastrales order (family: Lycopodaceae) of class, Hymenomycetes.

81. Sex in a saucer- Remnants of a cap of skull puffball on little bluestem-creepn juniper savanna in Little Missouri River Badlands. This was the fruiting body introduced in the first slide of skull puffball. The cap of this carpophore had disintegrated such that when the rounded top fragmented and blew away leaving the shallow base. There were still quite a few spores on the bottom of this remnant cap.

Theodore Roosevelt National Park (North Unit), McKinzie County, North Dakota. Late June.

82. Puffed over- Carpophore (fruiting body) of a fully ripe (spore-shedding) skullcap puffball on a little bluestem-creeping juniper savanna in Little Missouri River Badlands. This is the same carpophore (only upended) in the immediately preceding slide. The fruiting body--what most laymen think of as the mushroom "plant"--of a fungus consist of::1) cap (toadstool or parasol part) including gills, 2) stipe (the "stalk" or "stem" of the mushroom), 3) annulus (ring" on the stipe, and 4) volva (universal veil that can remain as a "cup" at base of stipe (near ground). Some of these organs, especially annulus and volva, are absent in some species as in most puffballs (like the species featured here).

More of these organs were presented in a photograph below of a more typical mushroom.

Theodore Roosevelt National Park (North Unit), McKinzie County, North Dakota. Late June.

83. Another fungus on northern plains range- Two pairs of fruiting bodies (carpophores) of meadow mushroom (Agaricus campestris) on a little bluestem-creeping juniper savanna (=creeping juniper dwarf shrubland, little bluestem-creeping juniper habitat type). According to standard descriptions in miscellaneous mushroom guidebooks this is one of the most widely distributed toadstool species in North America. Numerous of the Agaricus species (their fruiting bodies) closely resemble each other so that a spore print is the only way to get a definitive identification (a necesssity if and when carpophores are to be used as food, which was why this rangeman opted for roast beef or steak without mushrooms).

These top-down views were presented to show students that the cap (see immediately above) of the carpophore is divided into three portions for purposes of identification and description. These three zones of the cap are: 1) disc (apex or center of cap), 2 ) margin (outer edge of cap), and 3) limb (majority of cap that is between disc and margin). Note that margins of this species have indentations (ie. an indented cap).

The carpophores in the first slide were somewhat less ripe (mature) than those in the second slide. These meadow mushrooms grew within 20 to 25 steps of the skullcap puffballs described above. Grass species surrounding these fruiting bodies were western wheatgrass and plains reedgrass (little bluestem was the dominant grass species on this true prairie buffalo range.

Theodore Roosevelt National Park (North Unit), McKinzie County, North Dakota. Late June, maturing caprophores.

84. Morphology of carpophore- Lateral (side) view of a fruiting body of meadow mushroom (Agaricus campestris) that grew on a little bluestem-creeping juniper savanna in Little Missouri River Badlands. This carpophore was placed on the cap of a larger carpophore (these were the two fruiting bodies introduced in the second of the pair of slides presented immediately above). Easily distinguished parts of this fruiting body included: 1) remains of cup or universal veil, 2) stipe, 3) annulus on stipe, 4) gills on underside of cap, and 5) dorsal surface of cap on larger (still standing) carpophore. Features of the union of stipe and cap were also visible on the unearthed carpophore.

Although little blurestem was the dominant grass species on this little bluestem-creeping juniper savanna (creeping juniper-little bluestem habitat type) the shoots of the two principal grasses in closest proximity to the mushrooms were western wheatgrass and plains reedgrass.

Theodore Roosevelt National Park (North Unit), McKinzie County, North Dakota. Late June, maturing caprophores.

Vegetation classification and description note: the little bluestem-creeping juniper savanna treated above was probably most readily interpreted as a form of transitional vegetation between two ecological communities listed by the Montana Natural Heritage Program (2002): 1) little bluestem/plains muhly herbaceous vegetation and 2 ) creeping juniper/little bluestem dwarf shrubland.

85. Looks good enough to be bad- Landscape-scale view down the maw of Little Missouri River Badlands. Although tree-dominated vegetation consisting of Rocky Mountain juniper woodland and green ash groves were prominent in midground of this landscape the featured range plant community was the grassland of mixed prairie in foreground. This example of mixed prairie was included in both the True Prairie and Mixed Prairie chapters in order to show the continuum of various grassland range types extending from little bluestem-dominated grassland to little bluestem-creeping juniper savanna to this mixed prairie that included tallgrass, midgrass, and shortgrass species with often all of these rangeland cover types (or subtypes) contiguous with each other. This conglomerate grassland showed the continuity within these various grassland forms (= range types) as described by Barker and Whitman (1989) and in Shiflet (1994).

Mixed prairie vegetation seen here included prairie sandreed, which comprised the tallgrass component; little bluestem, western wheatgrass, sideoats grama, plains reedgrass, and plains muhly, which made up the midgrass that was the major component; and blue grama and Sandberg's bluegrass that together made up the shortgrass element. The main forb was wild or American licorice (Glycyrrhiza lepidota) which was shown below in "phytoquadrants" of this sward.

86. Good grass in badlands- Three landscape-scale photographs of grassland that developed on the relatively flat or level land immediately above the deeply eroded canyons of Little Missouri River Badlands. Grassland vegetation was a mosaic of patches of true and mixed prairie that comprised an overall mixed prairie. There were local overflow sites covered exclusively by western wheatgrass, shallow microsites dominated by blue grama, and isolated patches of prairie sandreed, but most grassland (most of that in foreground) was mixed prairie on which little bluestem was dominant with local botanical composition consisting variously of prairie sandreed, sideoats grama, plains reedgrass, and blue grama. The two most abundant forbs were wild or American licorice and stiff goldenrod. Shrubs were absent except for soapweed yucca on edges of the canyon.

Tree-dominated communities in background ranged from floodplain or river bottom forest of eastern cottonwood (Popuulus deltoids) through Rocky Mountain juniper woodland to groves of green ash.

87. Composition up close- Sward of the overall mixed prairie (with close resemblance to true prairie) introduced in the preceding three-slide set of Little Missouri River . Badlands grassland. In range vegetation of these two close-in photographs little bluestem was dominant and sideoats grama was associate species. Low, green forbs were wild or American licorice and stiff goldenrod.

Grassland vegetation at this scale--on this local microhabitat--was true prairie. This illustrated the fact that interpretation (hence classification and description) of a plant community is to some extent dependent on spatial (mapping) scale. Labeling such climax grassland as true versus mixed prairie depended on whether or not the beholder of this vegetation interpreted prairie sandreed and blue grama as plentiful (having sufficient cover) enough to qualify as contributing an ecologically meaningful tallgrass and shortgrass, respectively, component (layer) or, alternatively, if the major midgrasses were so overwhelming as to define the plant community irrespective of the limited presence of tallgrass and shortgrass species.

Note on location and organization: A mixed prairie variant of SRM 606 (Wheatgrass-Bluestem-Needlegrass) in the Northern Great Plains with a definite tallgrass component consisting of prairie sandreed (Calamolvilfa longifolia), prairie cordgrass (Spartina pectinata), and switchgrass (Panicum virgatum) along with little bluestem and without a Stipa species was presented and described in the Mixed Prairie chapter of Range Types of North America. There is a vast inter-regional (portions of Central and Northern Great Plains) mosaic of grassland consisting of tallgrass, true, and mixed prairies (complete with ecotones or transition zones among these) that extends from the Nebraska Sandhills northward through the Canadian Prairie Provinces and westward to the Rocky Mountains. Various range cover types within these units were orgainzed based on classification of vegetation and not as to geographic location or affinity.

Range Types of North America was organized according to historical and traditional treatments of range vegetation as to biomes, major formations, associations, and subdivisions of associations (ie. forest and rangeland cover types) and not as to continental regions based on physiographic provinces. Provinces and regions of North America were central to description and discussion of range dominance types, but orgainzation followed biomes and formations. Hence, range vegetation of the same area--even contiguous units of climax vegetation--were placed (are to be found) in different chapters. For example, grasslands and adjacent (bordering) forests with many of the same grasses were placed in separate chapters (= the former in one of the various chapters under Grassland; the latter in one--or more--chapters under Foorests).

Periodic notes on location, such as this one, were inserted at such strategic and confusing "organizational crossroads".

88. Grassland gradient- A "low knoll" (Barker and Whitman, 1989) in the Little Missouri River Badlands that sloped down to a level bottomland thereby forming a catena and/or a toposequence on which three distinctive range plant communities, corresponding to three recognized rangeland cover types (Shiflet, 1994), had developed. The Soil Science Society of America (2001) defined a catena ais "a sequence of soils of about the same age, derived from similar parent material, and occurring under similar climatic conditions, but having different characteristics due to variation in relief and in drainage" and the related term of toposequence as "a sequence of related soils that differ, one from the other, primarily because of topography as a soil-formation factor". Whether relief (topography) was the primary soil-forming factor responsible for diffenent soils and corresponding plant communities on this physiographic gradient was not known to this author, but soil drainage conditions and soil moisture content wa undoubtedly a major variable in the development of three distinctive climax range plant communities.

These three climax phytocommunities from level bottomland (foreground of both slides) to slightly increasing elevation just above base and lower sides of low knoll (midground of both slides) to upper sides and top of low knoll (background of both slides) three described rangeland cover types were: 1) needle-and-thread-Junegrass as co-dominants and thickspike wheatgrass as associate (SRM 607; Wheatgrass-Needlegrass), 2) western wheatgrass consociation (SRM 610: Wheatgrass), and 3) little bluestem-green needlegrass as co-dominants with sideoats grama as associate and some thickspikewheatgrass (variant of SRM 606: Wheatgrass-Bluestem-Needlegrass), respectively.

Theodore Roosevelt National Park (North Unit), McKenzie County, North Dakota. Late June, early estival aspect and society (peak standing crop and hard-dough stge for needle-and-thread and Junegrass). FRES No. 38 (Plains Grasslands Ecosystem). K-59 (Wheatgrass-Needlegrass). Mosaic of SRM types 606, 607, and 610. Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40). Northwestern Great Plains- Little Missouri Badlands Ecoregion 43b (Byce et al., Undated).

89. Local community of cool-season dominants- True prairie community co-dominated by needle-and-thread and Junegrass that developed on the bottomland portion of catena and/or toposequence in the Little Missouri River Badlands. This grassland phytocommunity was shown at its bottom edge or margin (lowest elevation) in the first photograph and at its upper margin (highest elevation) in the second photograph. At the upper elevational margin the needle-and-thread--Junegrass community met the lower elevtional edge of a western wheatgrass consociation (second photograph) forming a narrow transition zone (a microecotone) that was a mixture of the three dominants of these two grassland communities. Structure and composition of this needle-and-thread--Junegrass true prairie was more characteristic at the lower elevation margin (first photograph).

A little bluestem-green needlegrass community of true prairie had developed on the higher sides and top the short hill or what Barker and Whitman (1989) described as a low knoll (background of both slides).

90. Range vegetation in a true prairie drainage- Local shrub-dominated community in a low area with an ephemeral stream (wet-weather seepage to a light spring flow) in Little Missouri River Badlands. Range plants in this local corridor--a microsite--in true prairie included white sagebrush or Louisiana sagewort (Artemisia ludoviciana) and western snowberry (Symphoricarpos occitendalis) as co-dominants with western poison oak/ivy (Toxidodendron rydbergii= Rhus radicans,var. ryderbergii), Indian hemp (Apocynum sibiricum), and some species of beebalm or horse-mint (Monarda sp.) to round out this unique and localized range plant community which added further biodiversity to an already unique array and mosaic of true prairie climax vegetation (see again the preceding two sets of slides and captions).

This range community of dicotyledons was at the lowerest most margin of a needle-and-thread--Junegrass community (featured in the immediately preceding two slide-caption set) that was the low elevation portion of a toposequence of three grassland communities.

Theodore Roosevelt National Park (North Unit), McKenzie County, North Dakota. Late June, early estival aspect and society. No vegetational units for this local range plant community. Northwestern Great Plains- Little Missouri Badlands Ecoregion 43b (Byce et al., Undated).

91. Badlands true prairie- Smaller view of landscape of Little Missouri River Badlands with a ture prairie dominated in various consisting of a patchwork of local communities dominated variously by little bluestem, Junegrass, and/or needle-and-thread with numerous associates including Sandberg's bluegrass, blue grama, western wheatgrass, and thickspike wheatgrass. Little bluestem formed consociations that occurred as conspicuous patches or local populations that were readily distinguished by the tan-colored straw of last year's shoots as, for example, in immediate foreground of this photograph. Junegrass and needle-and-thread were found as either single-species consociations or, alternatively, as co-dominated (primarily just these two grass species) communities. Some forbs were common to all these communities which, again, formed a mosaic-pattern grassland.

These three dominants were midgrass species. Little bluestem grows to such reduced stature (as compared to little bluestem plants growing farther south and east in the central grasslands). Thus, this was textook true prairie and not mixed prairie of tallgrass and midgrass species. Blue grama and, evern more so, Sandberg's bluegrass were sparse enough in cover and plant density that, though present, they did not comprise a shortgrass componenet in this climax grassland.

In his--probably the--initial description of true prairie (Clements, 1920, ps. 121) labeled the true prairie as the Stipa-Koeleria association with S. comata, needle-and-thread, being dominant westward and S. spartea, porcupinegrass, being dominant eastward within this association. Clements (1920, p. 122) went on to specify that little bluestem was the normal associate of needle-and-thread/andor procupinegrass and Junegrass, the association-wide dominants.

This example of true prairie had developed on the outskirts of the Little Missouri River Badlands.

Theodore Rooseveldt National Park (South Unit), Billings County, North Dakota. Late June, late vernal aspect and society. FRES No. 38 (Plains Grasslands Ecosystem). K-59 (Wheatgrass-Needlegrass). Variant of SRM 606 (Wheatgrass-Bluestem-Needlegrass). Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40). Northwestern Great Plains- Little Missouri Badlands Ecoregion 43b (Byce et al., Undated).

92. Two communities of true prairie- Consociations of 1) needle-and-thread (foreground) and 2) little bluestem (background; easily discerned by straw or last year's dead shoots. Consociation is a climax plant community (sometimes relatively small in acreage or other dimensions) comprised of a single dominant species. Much of the true prairie in the Northern Great Plains exist as a mosaic of various consociations. Clements (1920, ps. 121-123) explained that in the true prairie little bluestem was the typical associate species with the dominants being 1) porcupinegrass and/or needle-and-thread and Junegrass. Needle-and-thread (Stipa comata) is the dominant Stipa species in the westward portion of true prairie, porcupinegrass (S. spartea) is dominant in the eastern part of true prairie, and the two are co-dominants in the central true prairie. Junegrass is a co-dominant across the true prairie (as an average or typical situation over the entire association); Junegrass is a three-way dominant where both Stipa species are dominants.

This grassland was on the outer margin of the Little Missouri River Badlands.

93. Splendor in the Badlands- True prairie consisting of consociations of Junegrass (most of the range vegetation in this slide), little bluestem (right background in this slide), and needle-and-thread (immediately preceding slide) that form a vegetational mosaic across this landscape that was on the outer margins of the Little Missouri River Badlands. Clements (1920, p. 121-123) explained that little bluestem was the typical associate species to needle-and-thread (or porcupinegrass) and Junegrass. In foreground of this photograph there were some plants of Sandberg's bluegrass that were part of a local population of this increaser grass, but overall the cover and density of this shortgrass species (and that of blue grama) was not enough to constitute a shortgrass component of this climax range vegetation.

There were no woody plants in the climax range plant community presented here. There were a few plants of forb species including prairie groundsel or prairie ragwort (Senecio plattensis) and downy Indian paintbrush (Castilleja sessilfolia). Specimens of both of these forb species were present in this scenic photograph.

94. Two cool-season domiants- Junegrass and needle-and-thread were co-dominants (and decreaser species) of the range plant community presented in these two "photoplots" taken on the border of the Little Missouri River Badlands. It was explained in the caption before the last (the immediately preceding) caption, the first caption that introduced this treatment of the needle-and-thread--Junegrass-little bluestem true prairie, that Junegrass and needle-and-thread and/or porcupinegrass are dominants of the Stipa-Koeleria association (Clements, 1920, ps. 121-123). These photographs showed that co-dominance of one of the Stipa species with Junegrass.

In the example presented here both of these cool-season, perennial festucoid grasses were at peak standing crop with caryopses ranging in maturity from milk to hard-dought phenological stages.

95. An uncommon consociation- Nested study plots (second slide was a smaller "photosample" within a larger near-landscape view) of a consociation of Junegrass that developed ou the border of the Little Missouri River Badlands. Junegrass was one of five grass species that Clements (1920, p.122) listed as forming consociations in true prairie showing importance of this midgrass species in true prairie. An even more revealing indication as to how important this pioneer plant ecologist regarded Junegrass was his naming of the true prairie the Stipa-Koeleria association (Clements, 1920, p.121), that is Stipa spartea and/or S. comata and Koeleria cristata were the overall or general (vs. local or restricted) dominants across the association .

Clements (1920, ps. 122) further indicated that of the five dominants that formed consociations in true prairie Junegrass was "exceptional" in being least apt to "occur as a pure dominant" and, furthermore, that Junegrass was the only one of these association-wide five dominants to exist strictly as a bunchgrass as the other four dominants are sod-formers (Clements, 1920, p. 123). These (and the next two) slides permitted students to see an uncommon--if not relatively restricrted- form of true prairie. Enjoy; compliments of God and the National Park Service.

96. Splendid cover- Sward of a consociation of Junegrass. Two progressively shorter camera-distance photographs of the structure and simple composition of a comparatively large stand of this native member of the Aveneae (oat tribe). There were a few plants of little bluestem in both of these "photoplots". (Viewers should note that little bluestem shoots are roughly the height as those of Junegrass showing, as was emphasized periodically throughout this section, that little bluestemis a midgrass in this portion of the Northern Great plains.) There were also some plants of prairie groundsel or prairie ragwort, a biennial or short-lived perennial forb.

Clements (1920, ps. 122, 123) explained that among the five dominants (all grass species) forming consociations within true prairie, the Stipa-Koeleria association, Junegrass was the dominant plant least likely to form consociations "as a pure dominant", and that Junegrass was the only dominant species which is strictly a bunchgrass and not a sod-forming grass.

Junegrass is also one of the most beautiful cool-season grasses of the great grasslands of North America. Furthermore, Junegrass is one of the most widely distributed grass species (has one of the largest species ranges) in North America being found from Quebec to British Columbia and south into Mexico.

The climax range vegetation seen in these and preceding (especially the immediately preceding two) photographs was relatively uncommon and certainly uncommonly beautiful. Aesthetics--of natural beauty in particular--is one of the multiple uses of range.

Location note: A green needlegrass-western wheatgrass cover type (SRM 607) was presented in the Grassland chapter entitled, Mixed Prairie, Northern Great Plains. Something of a case could be made for regarding SRM 607 as a form of true prairie, but a much stronger case can be presented for regarding that rangeland cover type as mixed prairie (See Clements, 1920, esp. 121-124, 135-138). The closeness and close proximity of true and mixed prairie in this range region was noted, and students promptly were referred to that chapter under the Grasslands heading in table of contents.

Western Wheatgrass True Prairie

Western wheatgrass (Agropyron smithii) is one of the most widely distributed midgrass species over much of the Great Plains. Its biological (species) range extends from (Barkworth et al., ). Western wheatgrass is also a species with almost as wide a distribution of scientific names (binomials). It was explained in the Names section of Introduction of Range Types that over course of the last quarter the genus of western wheatgrass century had changed numerous times including Elytrigia, Pascopyrum, and Elymus (besides Agropyron).

More pertinent to range vegetation is the situation or phenomenon that western wheatgrass occurs in a number of range plant communities including consociations (of nerly single-species stands) down to being a minor species in diverse grassland communities. Western wheatgrass occurs as a dominant in five rangeland cover types recognized by the Society for Range Management (Shiflet, 1994): SRM 606, SRM 607, SRM 608, SRM 609, and SRM 610 with this latter being "dominated almost exclusively by western wheatgrass" (Shiflet, 1994). Given that western wheatgrass is a midgrass SRM 610 (by definition more a population than a community) is true rather than mixed prairie.

In the seminal descriptions and comparisons of true and mixed prairies (Clements, 1920, p.122, 137) western wheatgrass and needle-and-thread were shown as the only tww species that formed consociatiosn in both of these grassland associations. Hence, distinction between true and mixed prairie on basis of species make-up can be tenuous and in the eyes of the beholding plant ecologist.For this reason samples of western wheatgrass-dominated grasslands were included in the two Grassland chapters, True Prairie and Mixed Prairie.

97. More like a field crop- Expanse of a western wheatgrass consociation that was in effect a nearly single-species stand (essentially a diverse population) of this member of the wheat or barley tribe (Hordeae or Tritaceae). This grassland vegetation developed on a seasonally wet habitat--Dense Clay range site--in a lowland in general area of the confluence of Little Missouri River and Bad River. It was a seasonal cattle range that appeared to have been grazed later in the previous year's grazing season. It had not been grazed during the current grass-growing season.

The primary soil of this lowland habitat was a gumb-like or heavy clay of the Swanboy (Swanboy clay) series

From descriptions of the climax grassland vegetation of this range site (Soil Conservation Service, 1975, p. 63; Soil Conservation Service, 1980, p. 92) western wheatgrass is the dominant and green needlegrass the associate species (roughly 60% and 20% of species composition, respectively, SCS, 1980, p. 92). A rough approximation of standing crop in the late boot to early spike-emergence stages of phenology would be that conservatively 9/10ths of biomass was of western wheatgrass. Overwhelming domination by western wheatgrass was probably due to previous overgrazing and/or some other disturbance given that green needlegrass declines and is replaced by western wheatgrass with overuse (Soil Conservation Service, 1975, p. 63). Green needlegrass was essentially absent from this range except for a few occasional plants (see the one-slide/caption unit below).

There were local (and very small) patches of buffalograss within which there were some plants of poverty sumpweed (Iva axillaris), an annual (and native) composite. The annual, Eurasian (now naturalized over much of North America) weed known as goatsbeard or salisfy (Tragapogon dubia) was also sparsely scattered throughout this otherwise single-species stand of grassland. Small patches of buffalograss likely reflected some form of disturbance (perhaps overgrazing) because the climax range vegetation lacks a shortgrass layer (Soil Conservation Service, 1975, p. 63). Plains prickly was the only woody species present and only as comparatively small plants (ie. not sprawling plants covering sizeable area) as was shown below in the last pair of photographs of this range plant community. There were also microsized (comparatively tiny) areas covered by Japanese brome or Japanese chess and/or Kentucky bluegrass, but these were incidential and limited to otherwise bare within more open spots in the sod of western wheatgrass.

Details of the plant community on this cattle range were presented immediately below. This pair of slides provided summary views of this range vegetation with emphasis on physiogonomy.

Fort Pierre National Grassland, Stanley County, South Dakota. Late June, vernal society with western wheatgrass at advanced boot to early spike-emergence phenological stages. FRES No. 38 (Plains Grasslands Ecosystem). K-59 (Wheatgrass-Needlegrass). Variant of SRM 607 (Wheatgrass-Needlegrass) degraded to SRM 610 or SRM 610 with potential component of green nedlegrass. Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40). Northwestern Great Plains- River Breaks 43c Ecoregion (Bryce et al., Undated).

98. Structure without much diversity- True prairie consisting almost exclusively of western wheatgrass in greater floodplain at confluence of Little Missouri River and Bad River in the glaciated Northern Great Plains. Water permeability was low on this Dense Clay range site so that it was a seasonally wet prairie. Green needlegrass is the associate species of the climax grassland, but past distubances (most likely overgrazing or even farming) resulted in nearly complete loss of green needlegrass from this particular range. The forb seen here was poverty sumpweed.

Both structure and species composition of the western wheatgrass consociation were apparent in these two photographs. There were patches of buffalograss with plants of plains pricklypear dispersed in these local (and small) areas dominated by shortgrass. There were even smaller microsized areas populated primarily by Japanese chess or Japanese brome and/or by Kentucky bluegrass. These little spots occurred on land where western wheatgrass shoots were of noticeably lower density. Green needlegrass had been almost extirpated from this specific range.

99. An open and uniform range turf- Features of the sward of a consociation of western wheatgrass in the confluence floodplain of the Little Missouri and Bad Rivers in the glaciated Northern Great Plains. Western wheatgrass is a strongly rhizomatous sod-forming grass so that on this Dense Clay range site almost all biomass was that of this species. Green needlegrass was the potential associate species in the climax vegetation of this range site (Soil Consrvation Service, 1975, p. 63) and its nearly complete absence spoke strongly of past abuse as green needlegrass is displaced by western wheatgrass with improper grazing (Soil Conservation Service, 1975, p. 63).

100. Structure and composition up close- Sample of western wheatgrass consociation that developed on the upper floodplain of converging Bad and Little Missouri Rivers. This "photoplot" provided viewers with a composite view of lowland true prairie in a comparatively high successional status. In addition to some young (immediate pre-anthesis) spikes of western wheatgrass there were some plants of the Japanese brome, an Eurasian annual that has naturalized across much of the North American mainland. Also in this photographic plot was one pre-bloom specimen of green needlegrass (cespitose plant smackdab in the center). This plant and a precious few others of green needlegrass attested to the potential presence of this species as the climax "second-in-command" (associate) for this Dense Clay range site (Soil Consrvation Service, 1975, p. 63). Past disturbances (farming, overgrazing, commercial traffic, etc. ?) had apparently eliminated green needlegrass from this true prairie range. Presence of this one plant of green needlegrass rendered this vegetational view as an unrepresentative and atypical example of the range plant community on this particular pasture. The slide was included to show that the potential vegetation was a western wheatgrass-green needlegrass climax.

101. Disturbance diversity- Local patch of buffalograss and plains pricklypear that developed due to spot grazing in a consociation of western wheatgrass in the vicinity of convergence of Bad River and Little Missouri Rivers. Spot grazing is the grazing pattern (and consequent vegetation) due to "repeated grazing of small areas while adjacent areas are lightly grazed or unused" (Kothman, 1974). Such erratic or "patchy" use is a sign of improper grazing management. Range animals should be properly distributed on the range so that the proper degree of use will be achieved over the entire range. Having some areas (regardless of size) overused (like this one) while other areas of the range are underutilized or not used at all is improper pasture management.

The invasive plains pricklypear was present on this range only in overgrazed patches such as viewers saw here. Viewers were also reminded that although proper distribution of range animals is essential for proper management of ranges, this uniform (even) distribution--hence, uniform grazing defoliation--in and of itself is not adequate for proper range management. Overgrazing across the entire range (as achieved by proper distribution of use) is far worse than localized spot-overgrazing like that seen here. Proper animal distribution must always be accompanied by proper degree of use, which is achieved through the correct stocking rate (proper number of animal units= animal numbers consistent with grazing or carrying capacity of the range). Overgrazing is longterm overuse (exceeding proper degree of use at any point time is overuse) that results in changes in species composition of the range plant community, lowered quantity/quality of forage soil erosion, etc. The ultimate result of overgrazing is a degraded or depleted range. Patches of buffalograss and plains pricklypear on this range site where the potential plant community was western wheatgrass and green needlegrass were small, localized areas of degraded grassland range. Overuse of small spots on this grassland over long periods of time resulted in spot overgrazing. Spots or small patches of this true prairie were converted into shortgrass prairie or, as more correctly described, shortgrass-cactus savanna. Grazing of this western wheatgrass-dominated range with even greater numbers of animals and/or grazing longer (or at the wrong season or stage of plant development) would have killed out even more wheatgrass which would have been replaced by buffalograss and plains pricklypear (ie. converted even more true prairie to a shortgrass-cactus savanna).

By the way, this is a textbook case where greater biodiversity was not good, not desirable. Increased diversity of plant species and of the range plant community was reached through localized overgrazing (spotgrazing) resulting in patches invaded by plants of lower successional rank or order (climax plant species replaced by seral species, including low-growing brush).

Directions to most of western wheatgrass range types- Most grasslands and savannas in which western wheatgrtass is a dominant, associate, or merely a major species falls under the category of mixed prairie. It is only range vegetation comprised nearly exclusively of western wheatgrass or of midgrass species and without shortgrass species, as other than sporadic individuals (ie. lacking a shortgrass layer or component), that "qualifies" as true prairie or true prairie-savanna. For this reason most range plant communities dominated (or, at least, defined) by western wheatgrass were more correctly included in the Grassland chapters devoted to mixed prairie (see especially the chapter, Mixed Prairie I).

Northern Mixed Prairie Savanna or Northern Mixed Grass-Shrub Steppe

Within the Northern Great Plains there are several range types (subtypes or variants) of grass-shrub savanna or shrub steppe. The grass component--the dominant and efeining layers--of these range plant communities are either mixed or true prairie, depending on major Gramineae species. The most common shrub(s) in this range vegetation is one or more species of sagebrush (Artemisia) which occur in pristine plant communities as relatively widely scattered, individual plants amid various combinations of tall-, mid-, and/or shortgrass species which comprise the predominant life or growth form. While the shrubs are conspicuous they are associate species (at best, aspect dominants).

The various forms of this shrub-grass savanna reflect a botanical affinity with the shrub-steppe climax of the Palouse Prairie which was interpreted by Weaver and Albertson (1956, ps. 340-341) as having extending across the Northern Great Plains to the Black Hills and Badlands of present-day Dakotas during portions of the Ice Ages when similar climates existed in the Interior Northwest and Great Plains province.

Several of these range types (and their variants) occurred contiguous with each other, and given that designation as to true versus mixed prairie was strained--if not arbitrary--distinction, this grassland vegetation was included both here as mixed prairie-shrub savanna as well as in the chapter, True Prairie.

102. A variant form of badlands savanna- In Little Missouri River Badlands a western wheatgrass-silver sagebrush savanna. This savanna vegetation had developed in a mosaic of (conterminous with) Rocky Mountain juniper (Juniperus scroplorum) woodland at slightly higher elevations at foot slopes of low hills and with other grassland communities at similar elevations on benches, swales, and bottomlands.

The second slide presented details of the silver sagebrush-western wheatgrass savannah at closer camera distance while the first slide gave viewers a landscape-scale perspective. Western snowberry (Smyphorocarpos occidentalis) was an associate shrub that commonly grew beside silver sagebrush and out in the sward of western wheatgrass. This spatial relationship was presented in the second of these slides.

With the adjacent development of Rocky Mountain juniper woodland and silver sagebrush savanna juniper naturally invaded (at least attempted invasion of) the grass-shrub savanna. This developmental phenomenon was described in detail in immediately succeeding slide-caption sets.

Theodore Roosevelt National Park (South Unit), Billings County, North Dakota. Late June, early estival aspect. There were no designations or descriptions of this savanna range vegetation. Instead it appeared that this range plant community was relegated to categorization as western wheatgrass-dominated grassland of the same general units presented above. FRES No. 38 (Plains Grasslands Ecosystem). K-59 (Wheatgrass-Needlegrass). SRM 610 (Wheatgrass). Cold Temperate Grassland 142, Plains Grassland 142.1 but no appropriate series in Brown et al. (1998). Northwestern Great Plains- Missouri Plateau Ecoregion 43 a (Bryce et al., Undated).

103. A Tale of Two Vegetational Cities-A Dickensonian version of two climax range types--one forest and one rangeland--in the Little Missouri River Badlands:1) Rocky Mountain juniper woodland on background slopes and 2) western wheatgrass consociation (essentially a single-species stand or population) on a bench that was overflow or swale site of plains grassland that at local and intermediate scale was a silver sagebrush-western wheaytgrass savanna. (This savanna vegetation was featured in the immediately preceding two-slide-catpion set.).

In the sample of range plant communities presented in these two photographs Rocky Mountain juniper was confined to its own woodland and was not in process of encroaching on (invading) the grassland/savanna range plant community. (This was in contrast to range vegetation described in the next slide-caption set.) Other woody species in the Rocky Mountain juniper-dominated woodland included the tree species, green ash and the shrub, skunkbush sumac (Rhus trilobata= R. aromatica). There were openings and smaller sporadic spots or patches of western wheatgrass within the juniper-dominated woodland. These western wheatgrass-dominated local areas often included little Indian ricegrass (Oryzopsis micrantha) as the associate species. Overall, for all practical purposes there was no herbaceous component--let alone, a layer--of herbaceous plants.within the Rocky Mountain juniper woodland. This was expecially the situation within interior of this tree-dominated plant community.

Theodore Roosevelt National Park (South Unit), Billings County, North Dakota. Late June, early estival aspect. Rocky Mountain juniper woodland was FRES No. 35 (Pinyon-Juniper Woodland Ecosystem), K-21 (Juniper-Pinyon Woodland), SAF 239 (Pinyon-Juniper; Rocky Mountain Juniper variant) and SRM 412 and 504 as aplied to Northern Great Plains, would be Pinyon-Juniper Series for Cold temperate Forest and Woodland 122 in Brown et al. (1998). Western wheatgrass grassland was FRES No. 38 (Plains Grasslands Ecosystem), K-59 (Wheatgrass-Needlegrass), SRM 610 (Wheatgrass), Cold Temperate Grassland 142, Plains Grassland 142.1 but no appropriate series in Brown et al. (1998) as it would be Wheatgrass Series under a Plains Shrub- Grassland. Northwestern Great Plains- Missouri Plateau Ecoregion 43 a (Bryce et al., Undated).

104. Ecroaching enemy-Seedlings and saplings of Rocky Mountain juniper invading a western wheatgrass consociation that developed on a benchlike swale in Little Missouri River Badlands. At another location within the badlands landscape a Rocky Mountain juniper woodland and western wheatgass-dominated grassland (a consociation) had developed contiguous with each other, but in this range vegetation the Rocky Mountain juniper had invaded and posed "a clear and present danger" to integrity and, in fact, continued persistence of the grassland. Fire exclusion by contemporary man over the last century had caused--at least contributed greatly to--brush invasion of virgin grassland ranges. This was one of numerous "footprints of the whiteman (in contrast to that of the American Indian whose frequent surface fires had preseved integrity of grasslands for several millenia.

Seen here was an early stage of the brush problem in the making.

105. Blessed be the fire; praise God for flames- Wild fire two years ago in this area of Little Missouri River Badlands killed various-sized plants of Rocky Mountain juniper that had invaded a grassland or grass-shrub savanna (depending on interpretation) dominated by western wheatgrass and green needlegrass with widely scattered plants of silver sagebrush and local patches (colonies) of western snowberry. There were small groves of young green ash on more favorable (more mesic especially) local sites. Wild fire had topkilled all plants of the two shrub and one decicuous (angiosperm) tree species, but all appeared to have resprouted. A number of years would be required before green ash attained the ages and sizes present before the fire.

By contrast the invasive--and nonsprouting--Rocky Mountain juniper were "graveyard dead". Hurray! Based on various features of the two major or most abundant grass species (standing crop or live shoot biomass, density, crown cover, general vigor, sexual reproduction) it was obvious that these climax dominants had benefitted tremendously (or at least not been adversely impacted) by the wildfire. Pre-burn measurements of these shoot features were not available, but prior experience and observation revealed that these characteristics or indications of plant growth and vigor were greater following than before the fire. Hurray, again!

Other than for small, local patches of Kentucky bluegrass there were essentially no other herbaceous plant species. Western wheatgrass and green needlegrass were clearly "monopolists".

The second of these slides was a closer-up view of silver sagebrush so as to feature the dominant climax shrub. These two photographs should be compared back to the two photographs of another western wheatgrass-silver sagebrush savanna in the Littel Missouri River Badlands that introduced this variant form of Northern Great Plains true or mixed prairie.

Theodore Roosevelt National Park (South Unit), Billings County, North Dakota. Late June, early estival aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-59 (Wheatgrass-Needlegrass). SRM 607 (Wheatgrass-Needlegrass). Cold Temperate Grassland 142, Plains Grassland 142.1 but no appropriate series in Brown et al. (1998). Northwestern Great Plains- Missouri Plateau Ecoregion 43 a (Bryce et al., Undated).

106. Release of the oppressed- Obvious increases in herbage yield (standing crop), shoot and plant density, basal and foliar cover, sexual reproduction, and greater plant vigor of western wheatgrass and green needlegrass were the bounty resulting from the flaming fingers of a wild fire two years ago on grassland or grass-shrub savanna in the Little Missouri River Badlands. Equally obvious--and more starkly conspicuous--were remains of dead Rocky Mountain juniper that had been killed by the fire. As in the case for almost all Juniperus species, J. scopulorum does not "grow back" (sprout or coppice) from shoot or stump. In Rocky Mountain juniper topkill is synomous with "complete kill" or "rootkill".

Invasion by Rocky Mountain juniper of grassland (such as those dominated by western wheatgrass) in landscapes like the Dakota Badlands is a widespread phenomenon in the absence of fire. Preceding examples of range vegetation in the Little Missouri River Badlands were presented in which Rocky Mountain juniper and western wheatgrass-doiminated grassland had developed continguously, but with grassland (interpreted variously as mixed or true prairie) being invaded by juniper in absence of fire.

Fire has been a natural part of Northern Great Plains landscapes and ecosystems, including those of badlands, for millenia. Fire is largely a component or outcome (or combination) of climate. Lightening-ignited fires are as natural a feature of climate as are droughts, blizzards, hail storms, etc. It could be argued that fuel (which completes the fire triangle with source of ignition and oxygen) is distinct from atmospheric factors. This is only partially the case because fuel sources are largely plant products (mostly cellulose) produced from such atmospheric components as precipitation, oxygen, nitrogen, temperature, and light. Of course fuel (vegetation) interacts with and greatly modifies the features of the atmosphere (eg. supplies much of the carbon dioxide in the air). Thus when certain range types, especially specific grasslands, shrublands, etc., are designated as "fire types" this merely highlights or signifies impacts of the apparently most immediate, obvious, or dramatic component of the regional climate. Ditto on "drought types". These range types are still classic Clementsian climaxes, now often labeled "climatic climaxes", with specific reference to that part of climate (fire, drought, etc.) that appears--largely by nature of its spectacular immediate affects--to be the most determinative factor for structure, composition, and function of these natural plant communities.

Anyway, fire killed the junipers that had invaded the grassland thereby largely restoring ecological integrity of the grassland or grass-shrub savanna.Thank God for fire, wild and natural or humanly prescribed and managed. Maintenance of natural grasslands and savannas requires fire (or some substitute that is generally in the form of ignited fossil fuels themselves an outcome of previous climates).

Resprouts of green ash (first slide) and silver sagebrush (second slide) showed that these woody species had through evolution (natural selection) adapted to fire the same as to other facets of climate such as winter temperatures, drought, and wind. By the way, fire is a defoliating agent the same as grazing animals. Resprouting species evolved to co-exist with fire the same as buffalo, deer, and jackrabbits. Rocky Mountain juniper has not undergone such evolution to recurrent fire. This is an example of the selective nature of "browsing" (defoliation) by fire. Take away that natural selective defoliator and vegetation that evolved with fire will change (as from grassland or savanna to woodland).

107. Death and renewed life from flames- In the Little Missouri River Badlands western wheatgrass and green needlegrass were the pyric beneficiaries from a wildfire two years ago on Northern Great Plains grassland dominated by these two climax fesutcoid grasses. This climax range vegetation could have been interpreted variously as: 1) mixed prairie due to some blue grama that might be interpreted as adequate for a meaningful shortgrass component, 2) true prairie dominated overwhelmingly by two midgrass species, or 3) mixed grass -shrub savanna on which silver sagebrush was the major climax woody species. Scattered plants of silver sagebrush, patches of western snowberry, and local groves of green ash were interpreted by this author as part of the climax range vegetation regardless of interpretation of the natural range plant community. On the other hand, by any of these three ecological perspectives fire was central to maintneance of the potential natural (climax) vegetation so that Rocky Mountain juniper was an invader, the establishment of which was due to unnatural fire suppression by past--if not present-- human action.

Wild fire results or impacts presented here (including previous slides) showed that the three major sprouting woody species (all angiosperms) were in the process of recovering whereas plants of the nonsprouting conifer (gymnosperm) were dead.Woody "skeletons" of Rocky Mountain juniper showed that these trees had been of comparatively large size with some of the larger ones apporaching sizes of the largest adults of juinper woodland in this same general area. Seedling-size sprouts (and perhaps seedlings as well) of we3stern snowberry, silver sagebrush, and green ash were distinct in the second of these slides.

Location note: Rocky Mountain juiper woodland (using examples from the Little Missouri River Badlands on Theodore Roosevelt National Park) were included with the two Woodland and Forest chapters, Juniper-Pinyon Woodland and Southern and Central Forests, in Range Types of North America.

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