True Prairie

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. 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.

3. 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.

5. 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.

6. 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).

10. 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.

11. 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.

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.

14. Autumn attire- Stand (first and second slide) and single cespitose plant (third 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.

15. Staying inside their shoots- Two 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.

16 . 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.

17A. 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.

17B. 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.

19. 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).

20. 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.

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).

21. 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).

22. 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.

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).

23. 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.

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).

24. 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).

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).

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.

25. 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).

26. 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.

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).

27. 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.

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).

28. 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.

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).

29. 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.

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).

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

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

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

31. 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.

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).

32. 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).

33. 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.

34. 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.

35. 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).

36. 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).

37. 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).

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).

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.

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.

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 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.

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).

39. 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).

40. 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.

41. 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.

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).

42. 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 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: 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.

43. 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).

44. 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).

45. 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).

46. 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).

47. 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).

48. 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).

49. 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).

50. 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).

51. 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.

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).

52. 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).

53. 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.

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).

54. 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.

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).

55. 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,

56. 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. .

57. 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.

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).

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.)

58. 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.

60. 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.

61. 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.

62. 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.

63. 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.

64. 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.

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).

65. 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.

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).

66. 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.

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).

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".

67. 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).

68. 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).

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).

69. 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).

70. 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).

71. 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.

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).

72. 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.

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).

73. 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.

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).

74. 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.

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).

75. 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.

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).

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.

76. 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).

77. 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.

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).

78. 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).

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).

79. 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.

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).

80. 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).

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).

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.

81. 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).

82. 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).

83. 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.

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).

84. 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).

85. 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).

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).

86. 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.

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).

87. Choice of cummunities- Two slightly views of typical landscape of Little Missouri River Badlands complete with a stretch of Little Missouri River in distant background. Range vegetation in these landscape scale photographs included: 1) floodplain (river bottom) forest dominated by eastern cottonwood; 2) small groves of green ash; 3) woodland of almost exclusively Rocky Mountain juniper; 4) grassland communities with varying proportions of western wheatgrass, grreen needlegrass, Junegrass, sideoats grama, little bluestem; 5) savannas of mixed grass-shrub species (especially silver sagebrush); 6) weed (mostly Eurasian grasses and forbs) patches across black-tailed prairie dog towns; and 7) populations of smooth bromegrass on disturbed areas such as road sides and ditches.

The range plant communities emphasized in these two views was native (climax) grassland and midgrass-silver sagebrush savanna seed in foregrounds. These range communities were shown and described in preceding and succeeding slides. Photographs of the overall Little Missouri River Badlands were inserted here to bring continuity to the "slide show" and to refresh viewers' perspectives of badlands range.

Theodore Roosevelt National Park (North Unit), McKinzie County, North Dakota. Late June, early estival aspect. Detailed enumeration of units for the range plant communities listed above were not provided in this caption. Northwestern Great Plains- Missouri Plateau Ecoregion 43 a (Bryce et al., Undated).

88. Dual lesson: Geology meets Ecology- An example of a slump in the Little Missouri River Badlands accompanied by a slide of National Park Service sign that explained the geologic phenomenon of slump formation. Range vegetation that developed by flow of water fro adjoining uplands was mixed grass-silver sagebrush savanna consisting of two major communities: western wheatgrass-silver sagebrush (foreground) and 2) western wheatgrass-green needlegrass-silver sagebrush (midground).

American Geological Insitute defined slump as: "a landslide characteritzed by a shearing and rotary movement of a generally independent mass of rock or earth along a curved slip surface (concave upward) and about an axis paralel to the slope from whidch it descends, and by backward tilting of the mass with respect to the slope so that the slump surface oftern exhitis a reversed slope facing uphill" ( Gary et al.,1972).

Closer-in (more detailed) views of these two range plant communities were presented immediately below.

Theodore Roosevelt National Park (North Unit), McKinzie County, North Dakota. Late June, early estival aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-59 (Wheatgrass-Needlegrass). Savanna variant of SRM 607 (Wheatgrass-Needlegrass) and 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).

89. Well-watered- General view (first slide) and close-in, detailed view (second slide) of a western wheatgrass-green needlegrass-silver sagebrush savanna in the Little Missouri River Badlands. Both co-dominant grasses grew in "nearly pure" local colonies such that there was a mosaic or patchwork of these two festucoid species. There were local patches of western snowberry (Symphoricarpos occidentalis) which qualified this shrub as an associate species.

This range vegetation was one of two plant communities that developed on gently sloping rangeland that was in a slump drainage immediately downslope from badlands uplands. Soil in this meso-scale range site was high enough in relative sand content which permitted green needlegrass to con-exist with western wheatgrass. This was in contrast to local habitats with higher clay-content soils that were single-grass species stands of western wheatgrass as shown in the next (succceeding) slide.

Theodore Roosevelt National Park (North Unit), McKinzie County, North Dakota. Late June, early estival aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-59 (Wheatgrass-Needlegrass). Savanna variant of 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).

90. Well-watered, but less diverse- Closer-in view of western wheatgrass-silver sagebrush savanna that formed on an overflow (swale) range site in Little Missouri River Badlands. There were some local colonies of western snowberry qualifing this shrub of the Caprifoliaceae as an associate species.

This range vegetation was one of two plant communities that developed on gently sloping rangeland that was in a slump drainage immediately downslope from badlands uplands. This local landscape was presented and described in the two slide-caption set before (immediately above) the last set. Soil in this local-scale range site was lower in sand content and higher in clay composition so that green needlegrass was not competitive with western wheatgrass which is the consumate swale-occupying/dominating native grass throughout much of the Northern Great Plains Region.

Theodore Roosevelt National Park (North Unit), McKinzie County, North Dakota. Late June, early estival aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-59 (Wheatgrass-Needlegrass). Savanna variant of 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).

91. Cannon ball rocks- Large exposed profile of land (probably several successive soil profiles) in Little Missouri River Badlands in which there were sandstone concretions of a size and roundnessas to suggest "cannon balls". The National Park Service sign provided an introduction to this phenomenon.

The American Geologic Instutute (Gary et al., 1972) defined concretions (of this form) as hard, compacted, typically rounded and spherelike aggregates of minerals formed by the precipitation process from liquid solutions in the pores of sedimentary (in this case) rock with a composition quite different from that of the original rock. Concretions are thus concentrations of the original rock and, often, some nucleus-forming object such as a fossil or bone along with cementing material. Sandstone concretions formed after initial deposition by cementation of sand grains

Range vegetation at base of this exposed (eroded) profile consisted almost entirely of western wheatgrass and silver sagebrush whereas green needltgrass was a co-dominant grass at distances farther from the eroded face (next slide).

Theodore Roosevelt National Park (North Unit), McKinzie County, North Dakota. Northwestern Great Plains- Missouri Plateau Ecoregion 43 a (Bryce et al., Undated).

92. When badlands get more water- A silver sagebrush mixed grass savanna with western wheatgrass and green needlegrass co-dominant on a flat near the raw, eroded face of a hill in the Little Missouri River Badlands. This range vegetation was farther away from the base of the erodec profile presented in the preceding three slide-caption set. Slightly sandier soil enabled green needlagrass to co-exist with western wheatgrass that is typically the sole dominant (often the sole species period) on heavier (more clayey) soils.

Theodore Roosevelt National Park (North Unit), McKinzie County, North Dakota. Late June, early estival aspect. Green needlegrass was in milk- to soft-dough grain stage. FRES No. 38 (Plains Grasslands Ecosystem). K-59 (Wheatgrass-Needlegrass). Savanna variant of 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).

93. True prairie through to the horizon- True prairie co-dominated by western wheatgrass and green needlegrass in the glaciated Northern Great Plains. An outstanding example of the climax (and pristine) form of wheatgrass-needlegrass rangeland cover type. Blue grama was an associate species and their were small, isolated patches of buffalograss on this range, but while these two shortgrass species present they were not abundant enough or with adequate foliar cover to comprise a shortgrass rcomponent (ie. this grassland vegetation was not even close to being mixed prairie).

There were isolated plants of Japanese chess (Bromus japonicus) and Kentucky bluegrass, but they would scarcely register as trace amounts. Forbs were almost non-existent on this climax grassland, but there were a few conspicuous plants of western yarrow (Achillea lanulosa= A. millefolium subsp. lanulosa). The most common or abundant forb was the naturalized Eurasian wild carrot or Queen Anne's lace (Daucus carota) while the most abundant dicotyledonus species was the suffrutescent composite, fringed sage or sagewort (Artemisia frigida). Plains pricklypear was present at a relative proportion (cover, density, etc.) that would be registered as Trace (given that there is not "sub-trace").

This range was being grazed by a commercial cow-calf herd at time of photographs. Management of all resources was as outstanding as this beautiful grassland that streatched from horizon to horizon.

Hughs County, South Dakota. Mid-June; late estival aspect, peak standing crop of green needlegrass. FRES No. 38 (Plains Grasslands Ecosystem). K-59 (Wheatgrass-Needlegrass). SRM 607 (Wheatgrass-Needlegrass). Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40). Northwestern Great Plains- Subhumid Pierre Shale Plains Ecoregion 43f (Bryce et al., Undated).

94. True to the horizon- Western wheatgrass-green needlegrass (co-dominants) true prairie in "mint condition" on the glaciated Northern Great Plains that extended to the far horizon. Blue grama was the warm season associate species, but it was not adequately abundant to add a shortgrass element to this grassland. This climax range vegetation was obviously of the wheatgrass-needlegrass dominance type (SRM 607) and not wheatgrass-gramagrass-needlegrass (SRM 608). There was also some buffalograss sporadically present in isolated patches that appeared to have been the result of widely spaced areas of heavy spot grazing (localized spots that had high degree of use) during the grazing history of this outstanding example of northern plains prairie range. Japanese chess and Kentucky bluegrass were present, but at cover and density that barely registered as the level of Trace.

Forbs were very sparse, but there were a few conspicuous plants western yarrow. The most abundant forb was the Eurasian biennial, wild carrot or Queen Anne's lace. The most abundant dicotyledon was fringed sage or sagewort, a suffrutescent species.

A number of sexual shoots of green needlegrass with caryopses in the soft-dough stage were prominently featured in the immediate foreground of all three of these slides.

This climax grassland was currently blessed with proper grazing management and all-around sound husbandry. It was being used as range for commercial cows and calves, and it was in Excellent range condition class. Rangemen responsible for the obviously extraordinary care given to this grassland were true to the best principles of grazing land management.

This Excellent condition range was a textbook example of the wheatgrass-needlegrass rangeland cover (dominance) type so the author felt it necessary to display three slides of such superb range vegetation (and superb range stewardship), the photographer's version of a three-rifle volley signifying great respect.

Hughs County, South Dakota. Mid-June; late estival aspect, peak standing crop of green needlegrass. FRES No. 38 (Plains Grasslands Ecosystem). K-59 (Wheatgrass-Needlegrass). SRM 607 (Wheatgrass-Needlegrass). Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40). Northwestern Great Plains- Subhumid Pierre Shale Plains Ecoregion 43f (Bryce et al., Undated).

95. Northern true prairie- Three landscape-scale views of pristine true prairie in the unglaciated Northern Great Plains. This obviously climax grassland was composed of numerous species of both bunchgrasses and sod-forming grass species along with widely scattered plants of Wyoming big sagebrush (Artemisia tridentata subsp. wyomingensis). Almost all grass species were decreasers including bluebunch wheatgrass, needle-and-thread, western wheatgrass, Junegrass, and blue grama all of which except blue grama were C3 cool-season species. There was lesser cover and shoot density of Sandberg's bluegrass which was an increaser on this range site. There was almost no cover of either cheatgrass or Japanese brome (this grassland was in "mint condition"). Numerous plants of threadleaf caric sedge (Carex filifolia) was present in local small areas of potential natural (climax) vegetation of this range. There were a few plants of soapweed or soapweed yucca (Yucca glauca). Also present at sparse cover was the subshrub or suffrutescent species known variously as fringed sage, fringed sagebrush, fringed wormwood, Arctic sagebrush, or prairie sagewort (Artemisia frigida). Locally common forbs included white beardtongue (Penstemon albidus), white milkwort (Polygala alba), smalleaf pussytoes (Antennariia parvifolia), slender milkvetch (Astragalus flexuosus), and pale evening-primrose (Oenothera albicaulis).

Blue grama is the major shortgrass species of this general region (Weaver and Albertson, 1956, passim ps. 327-343; Barker and Whitman, 1989) but on this and the subsequent grassland ranges described in this section blue grama was not present as a dominant and, only occasionally, as an associate. Such a conclusion was based on 1) personal observations by the current author and 2) the wheatgrass-needlegrass title (SRM 607) and description of this dominance type (Kuchler, 1964, p. 64; Shiflet, 1994). This climax grassland vegetation was regarded by this author true prairie not mixed prairie. In a somewhat differing interpretation (Barker and Whitman, 1989; Shiflet, 1994) regarded the wheatgrass-needlegrass type (SRM 607) as mixed prairie, but they along with Kuchler, 1964, ps. 64, 66) distinguished between the wheatgrass-needlegrass type (SRM 607) and the more xeric wheatgrass-grama-needlegrass type (SRM 608). Barker and Whitman (1989) and Shiflet (1994) specified that "the shortgrass component" was "notably more important" in this cover type than in the original climax vegetation. It must be stressed that the warm-season, eragrostoid blue grama was under-represented in the vernal society of this range. This was especially the case at stage of peak standing crop of cool-season species as shown in these (and subsequent) photographs. Such specification made, it was emphasized again that this was a wheatgrass/neeedlegrass-dominated range cover type with blue grama as no more than an associate species. (This discussion was continued in relation to the Palouse Prairie below.)

Either way, this was Northern Great Plains graassland as known by the Crow tribe (thought to be the first human group to live in this region).

Custer County, Montana. Mid-June, late vernal aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-57 (Grama-Needlegrass-Wheatgrass). SRM 607 (Wheatgrass-Needlegrass). Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40). Silty-Clayey range site complex, 10-14 inch precipitation zone, Eastern Sedimentary Plains (Ross and Hunter, 1976, p. 12). Northwestern Great Plains-Montana Central Grasslands Ecoregion 43n (Woods et al., 2002).

96. Harsh microhabitats- Local Breaks microsite range site within an overall Silty-Clayey range site complex of an Excellent condition range of the wheatgrass-needlegrass dominance type. The first photograph presented a locally restricted Breaks range site in left foreground that was surrounded by the deeper, more plant-favorable Silty-Clayey site. A nest of western harvester ant (Pogonomyrex occidentalis) indicated by the mound of excavated soil was at extreme margin of the Breaks range (right foreground). Low, flat rock outcrops characteristic of the Breaks habitat were present to the left of the ant nest (center foreground). Plant species more-or-less discernible in this larger-scale, first slide included bluebunch wheatgrass, western wheatgrass, needle-and-thread, Junegrass, Sandberg's bluegrass, and, among forbs or forblike plants, fringed sage. Shrubs included Wyoming big sagebrush and soapweed yucca. Visible in foreground was the cruciferous forb, comon pepperweed (Lepidium densiflorum).

The second photograph was a close-up view of a margin of this local Breaks range site adjoining the overall Silty-Clayey range site (left foreground and mid- to background). Of grass species listed for the first photograph the most abundant one in this slide was bluebunch wheatgrass (with lesser amounts of the other festucoid species) surrounding the rock outcrop of a local Breaks site. The major species growing on the outcrop spot were fringed sage, threadleaf caric sedge, and Sandberg's bluegrass. The showy forb on the outcrop was white beardtongue.

From the perspective of Landscape Ecology on this prairie the Breaks range site occurred as patches within a Silty-Clayey matrix.

Custer County, Montana. Mid-June, late vernal aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-57 (Grama-Needlegrass-Wheatgrass). SRM 607 (Wheatgrass-Needlegrass). Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40). Silty-Clayey range site complex, 10-14 inch precipitation zone, Eastern Sedimentary Plains (Ross and Hunter, 1976, p.12). Northwestern Great Plains-Montana Central Grasslands Ecoregion 43n (Woods et al., 2002).

97. God's own fenceline (two of His own fields)- The patchwork or mosaic of range vegetation on a wheatgrass-needlegrass true prairie was evident in this panning photograph. Local consociations of the dominant species grew in patterns of irregular shapes and various sizes as if there were small fields of different crops had been planted across the prairie, and indeed they were with God or Mother Nature the propagator. Range vegetation to the left of the foremost plant of Wyoming big sagebrush was a consociation of bluebunch wheatgrass while vegetation to the left of the big sagebrush plant was a consociation of western wheatgrass. Plants of Wyoming big sagebrush grew in both of these single species-dominated areas of climax range vegetation.

Physiogonomy, structure, and species composition of a major range cover type of natural grassland in the unglaciated western part of the Northern Great Plains.

Custer County, Montana. Mid-June, late vernal aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-57 (Grama-Needlegrass-Wheatgrass). SRM 607 (Wheatgrass-Needlegrass). Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40). Silty-Clayey range site complex, 10-14 inch precipitation zone, Eastern Sedimentary Plains (Ross and Hunter, 1976, p. 12). Northwestern Great Plains-Montana Central Grasslands Ecoregion 43n (Woods et al., 2002).

98. Native crop on one side of fence- Consociation of western wheatgrass within an overall wheatgrass-needlegrass true prairie in the sedimentary section of Northern Great Plains. The mosaic pattern of natural range vegetation formed by dominance of first one plant species and then another depending on range site and local microrelief is a characteristic feature of grasslands in this broad ecotone between climax communities of the Great Plains and those of the Rocky Mountain foothills. Such climax plant communities dominated by a single species are known in Clementsian terms as consociations. Consociation was defined by Allaby (1998) as a phytosociological term in long use by the Anglo-American school of Plant Ecology. Patchworks of consociations result in a mosaic of various climax grassland communities across landscapes of the Northern Gret Plains Region.

The first of these two photographs showed the western wheatgrass consociation at larger scale whereas the second photograph presented the sward of this single species-dominated climax vegetation at closer scale to show greater details of western wheatgrass shoots. This consociation occupied a large depression surrounded by slightly higher ground vegetated by a consociation of bluebunch wheatgrass. Needle-and-thread was the associate species on both Agropyron consociations. Western wheatgrass is best adapted to more mesic habitats such as those receiving runoff water (eg. depressions, swales, bottomlands). Bluebunch wheatgrass is a more xeric Agropyron species that is more abundant on upland sites. Western wheatgrass and bluebunch wheatgrass grow in side-by-side communities where microrelief consist of alternating low-lying and upland habitats.

Western wheatgrass is one of roughly four grass species that is a region-wide dominant (on various--though certainly not all--range sites across the Northern Plains). Needle-and-thread, blue grama, and green needlegrass (Stipa viridula) round out the big four dominants of this rangeland cover type (Kuchler, 1964, p. 66) on a regionwide basis. Bluebunch wheatgrass, one of the co-dominants of the Palouse Prairie, is frequently a dominant. in the western margins of the wheatgrass-needlegrass dominance type (SRM 607) treated in this section. (This was explained in greater detail in the immediately succeding caption.).

Western wheatgrass is the most widely distributed and generally most abundant Agropyron species in this zonal (regional) grassland. Western wheatgrass is also a dominant of mixed prairie dominance types such as the wheatgrass-grama-needlegrass cover type (SRM 606).

Custer County, Montana. Mid-June, late vernal aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-57 (Grama-Needlegrass-Wheatgrass). SRM 607 (Wheatgrass-Needlegrass). Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40). Silty-Clayey range site complex, 10-14 inch precipitation zone, Eastern Sedimentary Plains (Ross and Hunter, 1976, p. 12). Northwestern Great Plains-Montana Central Grasslands Ecoregion 43n (Woods et al., 2002).

99. Native crop on the other side of the fence- Consociation of bluebunch wheatgrass within an overall wheatgrass-needlegrass true prairie in the sedimentary section of Northern Great Plains. It was explained immediately above that on many ranges throughout this region microtopography consist of drier, upland environments situated beside mesic, lower-lying habitats such that consociations of bluebunch wheatgrass and those of western wheatgrass develop next to each other. This spatial organization of climax grassland vegetation was shown in the first of these two slides with a consociation of bluebunch wheatgrass in the foreground and one of western wheatgrass behind. Widely scattered plants of Wyoming big sagebrush grew in both consociations. Physiogonomy, structure, and coposition of this rangeland cover type were displayed in this larger scale slide.

The second of these slides showed structure and composition of the bluebunch wheatgrass consociation at closer distance (= smaller size "photoquadrant"). Some of the dried (sun-bleached or light-colored) sexual shoots of bluebunch wheatgrass were from the previous growing season.

Bluebunch wheatgrass is bettter adapted to more xeric habitats than is western wheatgrass. As such, on ranges with diverse microrelief, such as this one, bluebunch wheatgrass and western wheatgrass often grow adjacent to each other in single species-dominated patches of grrassland. Combined with such patches of dominant midgrasses were smaller microhabitats of Breaks range sites dominated by forbs and Sandberg's bluegrass, all of which resulted in a mosaic of climax vegetation characteristic of this rangeland cover type (Shiftlet, 1994)

Bluebunch wheatgrass is a relict species that has persisted on the Northern Great Plains over a geologic time span from when the Palouse Prairie extended eastward across the Northern Great Plains (Weaver and Albertson, 1956, ps.339-341). Presence of bluebunch wheatgrass, the overall co-dominant (along with Idaho fescue [Festuca idahoensis]) of the Palouse Prairie, was botanical proof of the affinity of climax grasslands on the western Northern Great Plains with those of the Palouse Prairie which currently is centered on the Pacific side of the Northern Rocky Mountains. Wheatgrass-needlegrass prairie in which bluebunch wheatgrass is a dominant species is one of the major range cover types in the broad ecotone between grasslands east of and those west of the Northern Rocky Mountains.

Custer County, Montana. Mid-June, late vernal aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-57 (Grama-Needlegrass-Wheatgrass). SRM 607 (Wheatgrass-Needlegrass). Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40). Silty-Clayey range site complex, 10-14 inch precipitation zone, Eastern Sedimentary Plains (Ross and Hunter, 1976, p. 12). Northwestern Great Plains-Montana Central Grasslands Ecoregion 43n (Woods et al., 2002).

100. Two compatriots- A local stand of western wheatgrass and needle-and-thread on a clayey phase of a range site of true prairie range in Excellent condition class that developed on uplands in the unglaciated Northern Great Plains. This "photoplot" presented tructure and composition of this local range plant community that was one of several consociations and various combinations in the vegetational mosaic of climax plains grassland.

Needle-and-thread was at soft-dough grain stage and peak standing crop. Western wheatgrass was in immediate prebloom stage with spikes still completely enclosed in the boot.

Custer County, Montana. Mid-June, late vernal aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-57 (Grama-Needlegrass-Wheatgrass). SRM 607 (Wheatgrass-Needlegrass). Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40). Silty-Clayey range site complex, 10-14 inch precipitation zone, Eastern Sedimentary Plains (Ross and Hunter, 1976, p. 12). Northwestern Great Plains-Montana Central Grasslands Ecoregion 43n (Woods et al., 2002).

101. An outlier of the Palouse Prairie-Two "photoquadrants" of progressively smaller size presented composition and structure of a wheatgrass-needlegrass true prairie range in Good condition class in the sedimentary section of Northern Great Plains. The first "photoplot" included three plants of Wyoming big sagebrush along with needle-and-thread, bluebunch wheatgrass, western wheatgrass (the "tri-dominants"), Junegrass, and threadleaf sedge. The second "photoplot" presented bluebunch wheatgrass (a fine specimen in left foreground), needle-and-thread (obvious with its florets filled with conspicuously awned caryopses), and western wheatgrass (not obvious in its prebloom-phenological stage). There were several small plants of fringed sage in foreground of second slide.

Blue grama was present, but not in cover even approaching that of needle-and-thread and the dominant wheatgrass species. Sparcity of blue grama was atypically representative in this vernal society. Nonetheless, blue grama (generally regarded as a shortgrass species) was not a major species and certainly not abundant enough to interpret this grassland as of the more droughty wheatgrass-grama-needlegrass type (SRM 608).

The wheatgrass-needlegrass prairie treated here and immediately below served as examples of the transition between grassalands east and west of the Northern Rocky Mountains. To some extent these wheatgrass-needlegrass ranges were "island outliers" of the Palouse Prairie.

Weaver and Albertson (1956, ps.339-341) described a wide ecotone or transition zone between grasslands of the Northern Great Plains and the Palouse Prairie. These workers hypothesized that at one time in the geologic past the Palouse Prairie extended from its present location in Pacific and Interior Northwestern North America across to the present-day Badlands of the Dakotas. Bluebunch wheatgrass and Sandberg's bluegrass persist as relict species in eastern portions of Northern Great Plains prairies in what once constituted the eastern margin of the Palouse Prairie (Weaver and Albertson, 1956, ps.340-341). Wyoming big sagebrush is the main component of a woody component showing the affinity of grasslands of Palouse Prairie (and the adjoining big sagebrush-bunchgrass savanna) with those of the grasslands to the east of the Northern Rocky Mountains. The western border of true prairie, main body of northern mixed prairie, and easternmost Palouse Prairie converge in the region of the Northern Great Plains.

Custer County, Montana. Mid-June, late vernal aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-57 (Grama-Needlegrass-Wheatgrass). SRM 607 (Wheatgrass-Needlegrass). Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40). Silty-Clayey range site complex, 10-14 inch precipitation zone, Eastern Sedimentary Plains (Ross and Hunter, 1976, p. 12). Northwestern Great Plains-Montana Central Grasslands Ecoregion 43n (Woods et al., 2002).

102. Details of the outlier- Representative "photosamples" of the sward of a wheatgrass-needlegrass prairie in the sedimentary Northern Great Plains. Species composition of range vegetation in the first photograph was mostly needle-and-thread, western wheatgrass, and bluebunch wheatgrass with a "smattering" of Sandberg's bluegrass. Plant life in the second photograph consisted mostly of western wheatgrass (local stand within a consociation of that midgrass species) with a fairly large plant of plains pricklypear for company. There were a few plants of Sandberg's bluegrass in this sample of range vegetation as well.

Custer County, Montana. Mid-June, late vernal aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-57 (Grama-Needlegrass-Wheatgrass). SRM 607 (Wheatgrass-Needlegrass). Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40). Silty-Clayey range site complex, 10-14 inch precipitation zone, Eastern Sedimentary Plains (Ross and Hunter, 1976, p. 12). Northwestern Great Plains-Montana Central Grasslands Ecoregion 43n (Woods et al., 2002).

103. Good, but not quite so nice-Another true prairie range of the wheatgrass-needlegrass type except that this was in Good range condition class versus the "in-mint-state" Excellent condition class range shown and described above. The range presented here was a different range site (though differening only slightly), but the main difference in species composition was most likely the result of heavier grazaing (apparently a grazing history that included some bouts of overgrazing). This range included relatively little bluebunch wheatgrass which had most likely been grazed out under past overgrazing (current degree of use was proper or, even, on the light side of proper use). There was much more cover of Sandberg's bluegrass, an increaser on this range type/site; Japanese chess, annual Eurasian invader, and sixweeeks fescue (Festuca octoflora= Vulpia octoflora), native annual. These two annuals were almost non-existant on the Excellent condition class wheatgrass-needlegrass range presented above. Students should note that Wyoming big sagebrush was of approximately the same cover and density on both the Excellent and Good condition ranges. Fringed sagebrush or fringed sage was also present on both the Good and Excellent condition ranges in roughly the same relative abundance and cover.

The greatest difference is species composition between these two range (and slightly different range sites) was presence of bluebunch wheatgrass on the former range versus high composition of thickspike wheatgrass (Agropyron dasystachyum) on this range and very sparse cover and density of thickspike wheatgrass on the Excellent condition range.

Other plant species on this Good condition range included the same forb species as on the Excellent condition range: white beardtongue, white milkwort, smalleaf pussytoes, slender milkvetch and pale evening-primrose. Additional forbs on this Good condition range included wavyleaf thistle (Cirsium undulatum), Missouri locoweed or Misssouri milkvetch (Astragalus missouriensis), purple milkvetch (A. agrestis= A. goniatus), shaggy fleabane (Erigeron pumilus), stiffstem flax (Linum rigidum), pale evening-primrose (Oenothera albicaluis), and pale agoseris or false dandelion (Agoseris glauca). Also present on this Good condition and not the Excellent condition range were numerous local colonies of lesser spikemoss or clubmoss (Selaginella densa), a pteridophytic (referring to vascular cryptogam) forb, along with lichen and field mushroom (Agaricus comprestis), two nonvascular cryptogams. (These cryptogams--along with meaning of cryptogam- were presented below.)

This range was in apparent upward range trend with recovery of the range plant community taking place through secondary succession following overgrazing. (This land had not been farmed, mined, etc. and instead was virgin--though degraded--sod,)

Garfield County, Montana. Mid-June, late vernal aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-57 (Grama-Needlegrass-Wheatgrass). SRM 607 (Wheatgrass-Needlegrass). Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40). Silty range site, 10-14 inch precipitation zone, Eastern Sedimentary Plains (Ross and Hunter, 1976, p. 11). Northwestern Great Plains-Montana Central Grasslands Ecoregion 43n (Woods et al., 2002).

104. Herbivores' home amid diverse herbage- Conical mound marking a nest of western harvester ant (Pogonomyrex occidentalis) on a wheatgrass-needlegrass true prairie range in Good condition class on unglaciated Northern Great Plains. Range plants included western wheatgrass, thickspike wheatgrass, needle-and-thread, Sandberg's bluegrass, white penstemon, wavyleaf thistle, Missouri locoweed, stiffstem flax, smallleaf pussytoes, and false dandelion or pale agoseris.

Garfield County, Montana. Mid-June, late vernal aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-57 (Grama-Needlegrass-Wheatgrass). SRM 607 (Wheatgrass-Needlegrass). Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40). Silty range site, 10-14 inch precipitation zone, Eastern Sedimentary Plains (Ross and Hunter, 1976, p. 11). Northwestern Great Plains-Montana Central Grasslands Ecoregion 43n (Woods et al., 2002).

105. Harvesters' garden- This local range plant community had developed along the outer perimeter of a "sacrifice area" that was made by western harvester ants around their underground "village" . Major range plant species within this disturbance vegetation of northern true prairie were threadleaf caric sedge (the large cspitose plant), Sandberg's bluegrass, needle-and-thread, shaggy fleabane, and stiffstem flax. The only woody plant species in the "photoquadrant" was plains pricklypear.

Garfield County, Montana. Mid-June, late vernal aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-57 (Grama-Needlegrass-Wheatgrass). SRM 607 (Wheatgrass-Needlegrass). Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40). Silty range site, 10-14 inch precipitation zone, Eastern Sedimentary Plains (Ross and Hunter, 1976, p. 11). Northwestern Great Plains-Montana Central Grasslands Ecoregion 43n (Woods et al., 2002).

106. More distant garden- At far edge of a localized plant community created by activiety of western harvester ants was this spreading, clonal plant of smallleaf pussytoes surrounded by needle-and-thread with a few plants of fringed sage. Out beyond this local "ant garden" major plants were western wheatgrass, thickspike wheatgrass, needle-and-thread, threadleaf caric sedge, Sandberg's bluegrass and a host of range forbs including white penstemon, wavyleaf thistle, Missouri locoweed, stiffstem flax, smallleaf pussytoes, false dandelion or pale agoseris, and lesser spikemoss or clubmoss.

Garfield County, Montana. Mid-June, late vernal aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-57 (Grama-Needlegrass-Wheatgrass). SRM 607 (Wheatgrass-Needlegrass). Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40). Silty range site, 10-14 inch precipitation zone, Eastern Sedimentary Plains (Ross and Hunter, 1976, p. 11). Northwestern Great Plains-Montana Central Grasslands Ecoregion 43n (Woods et al., 2002).

107. Shortgrass species on true prairie- Sandberg's bluegrass (Poa sandbergii= P. secunda) on a true prairie in Good range condition class in sedimentary Northern Great Plains. This "hale and hardy" specimen (two views of the same plant) approached the maximum size for this festucoid (pooid) native that has traditionally been regarded as a shortgrass species. There are some harsh or severe range sites on which Sandberg's bluegrass is a climax dominant, but in the grasslands of the Northern Great Plains such habitats are limited to extremely shallow soils, rock outcrops, and so on. On other range types,such as the channeled scablands (Bluegrass Scabland, SRM 106; SAhiflet, 1994), Sandberg's bluegrass dominates range sites and microsites that are too severe for midgrasses like Idaho fescue or bluebunch wheaqtgrass.

This robust individual that was used as example of for northern true prairie was growing on the range featured directly above. Sandberg's bluegrass was an increaser on this range where the potential natural (climax) grassland vegetation was wheatgrass-needlegrass (SRM 607). Presence of Sandberg's bluegrass as an important component, especially at local scale, of this range plant community was indicative of past disturbance (almost assuredly of overgrazing as this was otherwise virgin sod). Sandberg's bluegrasas was a key indicator species for this range. It was explained above that this Good condition class range was in apparent upward trend with recovery of grassland vegetation via secondary succession.

Sandberg's bluegrass is another one of those species over which controversy and confusion exist with regard to the taxonomically correct specific epithet. In fact this is confusion encompassing numerous Poa species (or alleged species) that are closely related (or are perceived as closely related), morphologically and/or genetically. early on Rhydberg (1922, ps. 81-82) recognized used the binomial P. sandbergii whereas Coulter and Nelson (1909, p. 73) showed the species P. Buckleyana with P. sandbergii and P. tenuifolia as synonyms. Hitchcock and Chase (1951, p. 134) used P. secunda as the scientific name for Sandberg bluegrass with P. sandbergii (the binomial used by Hitchcock's predeceessor, Vassey) shown as a synonym. Great Plains Flora Association (1986, p. 1214) went back or tetained P. sandbergii and interpreted plants represented by that binomial as integrading with P. caanbyi while the names P. buckleyana and P. secunda were regarded as having been "misapplied by various authors".

Arnov (1981) compared P. secunda of South America with P. sandbergii of North America and concluded that they were the same species with P. secunda being the correct name while P. sandbergii would then be "relegated to synonymy". Kellogg (1985) used various techniques, including transplant experiments and morphological evaluations, and concluded that many recognized (published) species in what she described as "the Poa secunda complex" were "all part of the single widespread species, P. secunda". Kellogg (1985) included six recognized Poa species in her "Poa secunda complex". Barkworth et al. (2007, 586-588) accepted these results (pretty hard to ignore transplant experiments) and split "the Poa secunda complex" of Kellogg (1985) into two subspecies. Skinner (2010, ps.254-267) followed suit and lumped four of the Kellogg (1985) six (plus one more) that are found in Wyoming as subspecies of P. secunda.

The specimen displayed in these two photographs was Poa secunda subsp. secunda.

Garfield County, Montana. Mid-June; soft-dough grain stage.

109. Out-foxed- Foxtail barley (Hordeum jubatum) on disturbed area in unglaciated Northern Great Plains. This native perennial bunchgrass prefers the "new land" of denudation or disturbance. It is in effect a weed, an ecological invader. Foxtail barley is utilized as forage, but its awns can cause mechanical injury to mouths and eyes of grazing animals, especially upon drying. These barbed awns can even penetrate the thin skin of sheep.

The old-reliable Range Plant Handbook (Forest Service, 1941, G74) treated this noxious species, including descriptions of mechanical injury to wildlife (elk in particular). The importance of foxtail barley as a weed--though also as a colonizer and facilitator of plant succession on waste places--was evident by its inclusion in the 200 species list on the International Range Plant Identification Contest sponsored by the Society for Range Management (Stubbendieck et al., 1992, 210-211).

Custer County, Montana. Mid-June; pre-anthesis stage of phenology.

110. Glistening fox tails- Dew-holding spikes of foxtail barley that was growing on a disturbed microsite in the Northern Great Plains. There are three spikelets per node of the rachis with the center spikelet fertile (perfect) and having one floret while the two lateral spikelets are sterile (rudimentary, neuter, or, rarely, staminate), with all spikelets having narrow, awned glumes (Great Plains Flora Association, 1986, p. 1185).

Custer County, Montana. Mid-June; pre-anthesis stage of phenology.

111. With the fringe down low- Two plants of fringed sage, firinged sagebrush, fringed wormwood, Arctic sagebrush, or prairie sagewort (Artemisia frigida) on pristine wheatgrass-needlegrass true prairie on the sedimentary Northrn Great Plains. This subshrub or suffrutescent species is a remarkable widely distributed species ranging from North America to Eurasia including the steppes of Mongolia. In North America is species range is from the Northwest Territories and Nova Scotia south into Mexico.

Fringed sagewort is not particularily palatable, but it is consumed by livestock and wildlife depending on feed conditions. Abundance of fringed sage has generally been interpreted as a result of overgrazing. Fringed sagewort is, however, a member of the climax true prairie community (especially on harsh microsites) as evidenced by its presence in the virgin vegetation of this beautiful, Excellent condition grassland.

Viewers should take note of the lichen (immediately to the top of the right fringed sage plant) on the soil surface of this rangeland. (More on the presence of this lichen species below.)

Custer County, Montana. Mid-June, eraly stages of annual growth.

112. Dainty though not frail legume- Slender milkvetch (Astragalus flexuosus) in the unglaciated Northern Great Plains. This is one of the smaller, least robust of the Astragalus species. This daity beauty was growing on a shallow rock outcrop in a pristine wheatgrass-needlegrass true prairie in southeastern Monatana. A. flexuosus was not listed as a poisonous range plant by Kingsbury (1964). (The plants of slender milkvetch are so small that it would probably difficult for lievstock to consume toxic quantities of it). Just another nice little range plant; it is a niodulated, nitrogen-fixing legume.

Custer County, Montana. Mid-June; peak-bloom stage of phenology.

113. Petite petals- Flowers of slender milkvetch with the three kinds of petals (one standard or banner, two wings, and two fused keels) discernable.

Custer County, Montana. Mid-June; peak-bloom stage of phenology.

114. A larger (and not so frail) legume- Missouri milkvetch (Astragalus missouriensis) growing on a wheatgrass-needlegrass true prairie range in Good condition class in the Northern Great plains. (This Good condition range was featured shortly above.) The first of these two slides presented the aboveground portion of this papilionaceous legume in its entireity whereas the second slide featured the sexual shoot bearing both inflorescence and legumes (these were shown in greaater detail in the next slide-caption set.

Garfield County, Montana. Mid-June; flowering to immature fruit phenological stages.

115. Flowers and fruits on the northern plains- Papilionaceous flowers and legumes of Missouri milkvetch. Structure and individual petals of corolla visible in both photographs. Characteristic curled distal portion of legumes were presented in the second of these two slides and in the second slide in the immediately preceding slide-caption set.

Good reference for Missouri milkvetch (as well as other Astragalus species in western North Amreica) was Hermann (1966, p. 56-57; overall, 35-79).

Garfield County, Montana. Mid-June; flowering to immature fruit phenological stages.

116. Purple on the plains- Purple milkvetch (Astragalus agrestis= A. goniatus) growing beside Sandberg's bluegrass on a severely degraded wheatgrass-needlegrass true prairie range in the sedimentary Northern Great Plains. A. agrestis was not listed as one of the toxic Astragalus species by Kingsbury (1964). A agrestis was treated as synonymous with A. dasyglottis is Hermann (1966, p. 75-76) who described it as a "valuable species" often reported to be "highly palatable to livestock".

Garfield County, Montana. Mid-June; peak-bloom phenological stage.

117. Leaves and inflorescence (first slide) and details of papilionaceous flowers in head (second slide) of purple milkvetch on a depleted true paririe range. Leaves beside the head in second slide were those of Sandberg's bluegrass.

Garfield County, Montana. Mid-June; peak-bloom phenological stage.

118. Really a showy one- Two-groove milkvetch or two-groove poisonvetch (Astragalus bisulctus) on ungrazed range in the unglaciated Northern Great Plains. Even the most case-hardened, non-asthetically inclinded observers would have to grant that this is one of the most strikingly beautiful of all range fobrs in the immense Great Plains Region. This species is one of the more common and widespread Astragalus species that grows on seleniferous soils (Hermann (1966, p.38). In fact, A. bisulcatus is an obligate selenium accumulator and is therefore an indicator species of seleniferous soils (Kingsbury, 1964, ps. 45, 48, 306).

Crook County, Wyoming. Mid June- peak-bloom phenological stage.

119. Showy up close- Upper sexual shoots (first slide) and a spike-like raceme (second slide) of two-grove milkvetch or two-groove poisonvetch in the sedimentary Northern Great Plains.

Crook County, Wyoming. Mid June- peak-bloom phenological stage.

120. Crazy on degraded true prairie- Lambert crazyweed or Lambert's locoweed (Oxytropis lambertii) growing on a depleted mixed prairie. Lambert crazyweed is a widely distributed species throughout the Wester Range Region

This was the first species described and pictured (twice and in color) in Stock-Poisoning Plants of the Range (Marsh, 1918).It was in the colored frontispeace as a principal stock-poisoning plant in Range and Pasture Management (Sampson, 1923), the first textbook in Range Management. Durrell and Newsom (1936) described poisoning by this species (known at that time as Aragallus lambertii) on Colorado ranges. Of course O. lambertii was in Kingsbury (1964, ps. 307-311) and, with expanded coverage, in Burrows and Tyrl (2001, ps. 514, 594-599 passim), the current "bible" of toxic North American plants.

Emmon County, North Dakota. Late June; peak-bloom phenological stage.

121. Crazy papilionaceous- Details of corollas of the papilionaceous legume, Lambert's crazyweed or lambert locoweed, that was growing on depleted rangeland in the Northern Mixed Prairie. The flowers of this highly toxic species are unarguably showy.

Emmon County, North Dakota. Late June; peak-bloom phenological stage.

122. Not all is silver- Silver or white locoweed (Oxytropis sericea) growing on a severely overgrazed ture prairie. The potential natural (climax) vegetation for this range was grassland dominated by western wheatgrass, bluebunch wheatgrass, and neeedle-and-thread (a form of the wheetgrass-needlegrss type). Instead it was a grazing disclimax (disturbance climax) comprised mostly of Sandber'gs bluegrass, Japanese chess, downy brome or cheatgrass, and Wyoming big sagebrush. This woefully depleted range was shown and described below (same section of this chapater), but examples of silver locoweed growing on that overgrazed range were place here for inclusion with other herbaceous legumes of tribe, Galegeae.

O. sericea is one of the "classiest" of the poisonous range plants: "classy" as in showy or stuningly attractive and as a classic or textbook poisonous range plant. Silky crazyweed is one of several Oxytropis and Astragalus species regarded by Kingsbury (1964, ps. 307, 309) as "true locoweeds" (ie. produce typical symptoms of loco poisoning). This species usually made it into the standard texts of poisonous plants (eg. Cheeke and Shull, 1985, ps. 143, 146). In their poisonous plant compendium Burrows and Tyrl (2003, ps. 594-599) covered Oxytropis species, including O. sericea. (Incidentially this magnificant reference is highly recommended to all true rangemen. It should be a shelf companion to Kingsbury (1964).

Garfield County, Montana. Late June; peak-bloom stage.

123. Silver wings and keels- Papilionaceous flowers of silver or white locoweed produced on a plant happily at home on a severely overgrazed true prairie that was converted to a disclimax of Sandberg's bluegrass, Japanese chess, and Wyoming big sagebrush.

The papilionaceous flower has a corolla consisting of five petals": 1) one banner or standard (the largest andhindmost petal) to the front and side of which of which are 2) two wings, and at the foremost part of the corolla 3) two fused petals forming the keel.These three distinct forms of petal can be seen in this photograph (if the viewer looks closely enough).

Garfield County, Montana. Late June.

124. The cat's whiskers feet- Small-(little)- leaf pussytoes (Antennariia parvifolia) on a wheatgrass-needlegrass true prairie range in Good condition class on sedimentary Northern Great plains. This composite of the tribe was accompanied by lesser clubmoss or lesser spikerush and Sandberg's bluegrass. Other plant species adjoining this local colony of pussytoes included western wheatgrass, thickspike wheatgrass, needle-and-thread, and a "pasel" of range frobs including white penstemon, smallleaf pussytoes, stiffstem flax, wavyleaf thistle, Missouri locoweed, false dandelion or pale agoseris, and lesser spikemoss or clubmoss.

Garfield County, Montana. Mid-June.

125. Girls' night (and day) out- Closer view of the little-leaf pussytoes plant featured in the immediately preceding slide. This was a local colony (clonal group) of A. parvifolia made up of numerous small, circular units (modules) arising from (off of) stolons. This form of asexual reproduction is vegetative (propagation from shoots; more specifically, from meristematic tissue via adventituous shoot/root development). This is the same fundamental process as planting "sprigs", "runners", "eyes" (buds of tubers), etc. of agronomic (eg. forage grasses) and horticultural crops. Each or, at least, most of these clones or modular units ("daughter" or "sister plants") was also producing flowering shoots (tiller-like stems with inflorescences).

Antennaria species are dioecious. These flowering shoots--hence, the clonal units or modules--were all female. They were more than likely all clones of the same plant, the same genetic individual or genotype, that developed by the process of forming or growing new plant modules ("sister" or "daughter plants") from stolons. This developmental process (vegetative propagation) repeated successively resulted in the mat- or carpet-like phenotype seen here. Each of these phenotypic modules is a clone or ramet of the original "mother" plant which was the genet, the original genotype.

The flowering shoots of clonal or modular units were shown in greater detail in the succeeding pair of slides-- along with the "rest of the story".

Garfield County, Montana. Mid-June; full-bloom phenological stage.

126. Independent little range winches- Pistillate flowering shoots (first slide) and pistillate inflorescences (second slide) arising from a clonal mat of female modules or ramets ("daughter plants") of little-leaf pussytoes. This mono-genotypic (clonal) plant was part of the vegetation of a wheatgrass-needlegrass prairie range in Good condition class in the unglaciated Northern Great Plains.

Little-leaf pussytoes grows variously as sexual, asexual, and intermediate populations with sexual and intermediate populations found in southern parts of this species range (Bayer and Stebbins, 1987, p. 317). Although A. parvifolia is dioecious, male plants are comparatively rare in more northern parts of North America. In this northern region little-leaf pussytoes occurs as asexually reproducing female plants and populations. This is the condition designated as gynoecious, plants produce only female flowers so that seed--and not pollen--is the result from this asexual reproduction. Seed production without pollination is the phenomenon known as apomixis. In northern portions of its biological range A. parvifolia exist as female apomicts or, same thing, apomictic females.

Apomixis is another form of asexual reproduction. It is asexual seed production. This form of asexual propagation differs from asexual reproduction via vegetative multiplication (as when new plant modules arise and develop from stolons). Thus it is that female plants (and aggregate populations) of little-leaf pussytoes rely on two forms of asexual regeneration for perpetuation of their race, genotype, ecotype, (or whatever it is): 1) vegetative (stolons with vertical pistillate shoots) and 2) apomixis (fruit and fertile seed formtion).

It is important to realize that "new plants" of gynoecious (= all female "plants") little-leaf pussytoes produced from apomictic seed are also clones or ramets, "carbon-copies" of the original genotype or the genet. Whether these herbaceous little gals arise from apomictic seed off of gynoecious (pistillate) flowers or from vegetative modules off of stolons they are the same genetic individual (genotype) as great, great, great, great, great, ..........grandmaw. And there was no great, great, great, great, great, ..... grandpaw, at least not of this asexual female (gynoecious) genotype. Males not needed; males need not apply.

Garfield County, Montana. Mid-June; full-bloom phenological stage.

127. Ain't pussy footin' around- Local mat of small-leaf pussytoes (Antennaria parvifolia= A. parviflora) on a range near southern edge of Nebraska Sandhills. This plant was included here to complete the sequence of reproduction in this intriguing range plant. Modular units of this clonal (stoloniferous), perennial composite were caught by the Nikon at peak "harvest time" as they were sending forth their achenes each equipped with a tiny pappus. Pappus refers to a modified perianth that forms a crown on the tip of the achene (McGregor et all., 1986, p. 1324). This little composite of tribe, Inuleae, was using two forms of reproduction to retain its place in Nebraska Sandhills tallgrass prairie.

This story of reproduction in this short-statured composite becomes even more intriguing with "in-depth analysis". The Antenaria species are dioecious, but staminate plants of A. parvifolia are apparently quite rare over much of the Great Plains Region (McGregor et al, 1986, p. 861). Instead, fruit-production in this species in more northern parts of its range is predominately by apomixis (Bayer and Stebbins, 1987, p. 317). Thus even with seed production there is no recombination of germ plasm so that offspring arising from apomictic seed (inside achenes) would be clonal the same as daughter plants (modules or clonal units) coming off of stolons. The two forms of propagation--one vegetative; the other, seed production--in littleleaf pussytoes were both asexual reproduction.

I told ya'll this grassland gal wasn't pussyfooting around!

Lincoln County, Nebraska. Mid-June; fruit-shatter phenological stage.

Ending Range Ecology lesson from Miss Pussytoes: From the perspective of Evolutionary Ecology and "foraging strategy" (resource allocation in context of natural selection) there is one "terminal" lesson or "morale" to the tale of the gynoecious (asexual female) Antennaria parvifolia (= A. parviflora). Her asexual modes of reproduction (propagation by means of vegetative and apomictic seed) is an evolutionary deadend. The genotype cannot be changed. It "breeds true to type" every time and thereby insures maintenance of the current superior germ plasm, but it (the gynoecious form of the species) cannot be improved. Current populations of asexual female A. parviflora cannot genetically become better adapt to current habitat or, perhaps more importantly, to a changed (or changing) habitat. There is no process for genetic recombination and, therefore, there cannot be genetic improvement (no adaptation by means of natural selection) to a changing environment. Instead, all adaptation to a changing (changed) habitat must come through existing opportunities for change in phenotype. This is phenotypic plasticity, and the limits to phenotypic plasticity are the limits to survival of the presently existing plant (= current population of clones).

Survival (survival of the fittest) always comes down to one thing--and one thing only--successful reproduction in the species' habitat; a habitat that, as Palentology clearly shows, is going to change (if it is not always changing). Continued survival of clonal-female littleleaf pussytoes depends entirely on her present genotype being versatile enough to allow continued reproduction in the dynamic equilibrium of her grassland environment, the ebb and flow of the range ecosystem. Asexual reproduction is an evolutionary deadend.

The Eagles singing group said it in Lying Eyes: "...every form of refuge has its price"

128. Standing stiff- Single plant of stiff or stiffstem flax (Linum rigidum) on Northern Great Plains true prairie. Whole-shoot portion of plant (first slide) and upper part of shoot (second slide) on a wheatgrass-needlegrass range in Good condition class. This annual has a large biological range across the Western Range extending from the Canadian Prairie Provinces south into the prairies of central Texas and east to the former prairies of Illinois.

A plant of fringed sage, fringed sagebrush, or was growing immediately beside the stiff flax.

Garfield County, Montana. Mid-June; full-bloom stage.

129. Pair of pretties- Two inflorescences of stiff or stiffstem flax with their five two-toned petals. These showy flowers were on the shoot shown in the immediately preceding slide.

Garfield County, Montana. Mid-June; full-bloom stage.

130. Pale pretty on true prairie- Pale or prairie evening-primrose (Oenothera albicaluis) on wheatgrass-needlegrass true prairie that was in Good range condition class on Northern Great Plains (unglaciated portion). This short (no more than five inches in height) little rascal had as some of its forb "phyto-neighbors": stiffstem flax, little--leaf pussytoes, Missouri milkvetch, purple milkvetch, shaggy fleabane, and pale agoseris or false dandelion. (This Good condition range was described above as introduction to this section on forbs of Northern True Prairie.)

This little annual also occurs widely in mixed prairie, especially on overgrazed grassland such as that of prairie dog towns. One such example of pale evening-primrose on a "dog town" was presented in the Central Great Plains Mixed Prairie section of Mixed Prairie-I (under Grasslands in table of contents).

Garfield County, Montana. Mid-June; full-bloom stage.

131. A really white one- White milkwort (Polygala alba) on wheatgrass-needlegrass true prairie in the Northern Great Plains. These were two specimens (each slide was of a different plant) were growing on the pristine range (Excellent range condition class) described above that was interpreted as an outlier of the Palouse Prairie. P. alba is a deep tap-rooted perennial. This condition resembles the "foraging strategy" of many alpine-plant species: have most biomass belowground and flower most profusely only during exteremely favorable years (like the record wet year when this delightful find was captured by the Nikon FM). Even under best of growing conditions a body's got to be tough to get through a Montana winter-- or a Texas summer--and this species is extremely adaptable having a species range from Alberta and Saskatchewan south into Mexico.

Custer County, Montana. Mid-June; first plant (first slide) was at peak-bloom phenological stage whereas the second plant (second slide) was at early to mid-bloom stages (individual inflorescences at different stages of flowering).

132. Coming to her milk- Inflorescences (spike-like racemes) of white milkwort at early to mid-bloom stages. These flowering shoots were on the second plant (second slide) presented in the above slide-caption set (ie. in the immediately preceding photograph). Polygala species are in the Polygalaceae, milkwort family.

Custer County, Montana. Mid-June; early to mid-flowering stages.

and

and

133. Palefaces on former Crow hunting grounds- Paired photographs of two plants (first slide of both pairs) and upper potions of their sexual shoots or inflorescences (second slide of both pairs) of white beardtongue (Penstemon albidus) on wheatgrass-needlegrass ture prairie in unglaciated Northrn Great Plains. The first two shoot and flowers (first pair or upper pair of slides) were on the Good condition range presented and described above. This specimen was growing on relative deep soil and had numerous neighboring forb species (many of these were shown previously in this section). The second two shoots and flowers (second pair or lower pair of slides) were on the pristine climax grassland of bluebunch wheatgrass, needle-and-thread, western wheatgrass, and Junegrass featured above. This specimen was growing on an extremely shallow soil of a rock outcrop and did not have much by way of "forbaceous" neighbors.

Plants of white beardtongue are some of the smalleset individuals of the numerous Penstemon species growing across the vast Western Range Region. A standard reference for the Penstemon species in this region was Rickett (1973, ps. 592-609).

The Crow tribe had once possessed this land when they were forced westerward by warring Sioux who in turn had been forced west by the predominately Anglo-Americans (palefaces with the great white father living in the White House).

134. Details in white (mostly)- Individual flowers on portion of flowering shoot of white beardtongue on wheatgrass-needlegrass true prairie range in the sedimentary Northern Great Plains. These flowers were on a plant on the pristine range of bluebunch wheatgrass, needle-and-thread, western wheatgrass, and Junegrass shown and described above. By standards of many beardtongue species plants of white beardtongue are little guys. They are eye-catching, especially for being so small.

Custer County, Montana. Mid-June; full-bloom phenological stage.

135. It ain't all grass (or even grass and forbs) on grassland- Local stand of lesser spikemoss escorted by a fruiting body caprophore of common field mushroom (Agaricus compestris) and some species of lichen on a wheatgrass-needlegrass prairie in Good condition class on the Northern Great Plains. There were a few leaves of Sandberg's bluegrass and needle-and-thread visible in this photograph.

Garfield County, Montana. Mid-June, late vernal aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-57 (Grama-Needlegrass-Wheatgrass). SRM 607 (Wheatgrass-Needlegrass). Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40). Silty range site, 10-14 precipitation zone, Eastern Sedimentary Plains (Ross and Hunter, 1976, p. 11). Northwestern Great Plains-Montana Central Grasslands Ecoregion 43n (Woods et al., 2002).

136. Cryptogams on wheatgrass-needlegrass range- Local colony of lesser spikemoss (Selaginella densa) with some plants of a lichen on a western wheatgrass, thickspike wheatgrass, needle-and-thread-Sandberg's bluegrass prairie range on the Northern Great Plains. This natural pasture was in Good range condition class. Some culms of Sandberg's bluegrass were present.

Cryptogams are plants that reproduce by spores or similar propagules other than seeds. Vascular cryptogams are pteridophytes, nonflowering spore-producing vascular plants that have alternation of (and morphologiclly distinct) generations. This is the Pteridophyta division of the Plantae (plant kingdom) of the classical Linean hierarchial scheme. Included in Pteriodphyta is Selaginellaceae, the family of clubmosses or spikemosses with species like Selaginella densa shown here. Lichens are included in the nonvscular cryptogams of classical or Linean taxonomy (the format used throughout Range Types of North America). In the Linean scheme nonvascular cryptophytes such as lichens were regarded as thallophytes, those plants in the subkingdom, Thallobionta. Thallophytes (lichens, fungi, most algae are those plants which are not differentiated into roots or shoots. Lichens are composite plants that are g comprised of a species of fungus and a species of alga.

Garfield County, Montana. Mid-June.

137. Spiked on the plains- Lesser spikemoss (Selaginella densa) on a wheatgrass-needlegrass prairie in Good range condition class on the Northern Great Plains. Closeup views of the plant shown in the preceding slides. It appeared from literature searches that little has been written about the basic botany or biology of this species. It was perhaps revealing that one of the best discussions of the basic biology of Selaginella spp. were in older Botany text (Eames, 1936, ps. 31-47; Holman and Robbins, 1946, ps. 523-530).

Garfield County, Montana. Mid-June.

138. Fire on a rock- Desert fire-dot lichen (Caloptaca trachyphylla) growing on a large slab of sandstone of which there nare umerous varied forms in Garfield County, Montana (Natural Resources Consrvation Service, 2007). This large rock with its interesting lichen was on a wheatgrass-needlegrass true prairie in the Northern Great Plains (unglaciated portion). A good discussion of this lichen species was in the extraordinary reference, Lichens of North America (Brodo and Sharnoff, 2001).

Lichen, the symbiotic (mutualistic) growing together of algae and fungi, is part of the plant life of this grassland ecosystem. It is part of the range vegetation the same as the grasses and sagebrush. This dramatically colorful example was included as reminder of that fact to rangemen, newcomers to oldtimers.

Garfield County, Montana. Mid-June.

139. Now for one in Fair condition- A wheatgrass-needlegrass range on the unglaciated Northern Great Plains degraded to Fair range condition class. Past overgrazing had reduced relative cover of western wheatgrass, bluebunch wheatgrass, thickspike wheatgrass, and needle-and-thread and an increase in Sandberg's bluegrass (current dominant), threadleaf caric sedge, Japanese chess among herbaceous species combined with a large increase in Wyoming big sagebrush, the only woody species that was common on this range.

Degration of (the process of retrogression on) this range had been in the historic past. Current degree of use was on the low (light) end of moderate. This range was a good example of range deterioration due to past--not current--grazing practices, in this instance overgrazing. In fact, given present degree of use (a reflection of stocking rate), vigor and reproduction of range plants, and condition of soil surface (including littter) there was apparent range improvement. This rangeman concluded--tentatively--that range trend was up on this pasture.

These two slides presented to viewers a general or composite view of the general landscape and relief--the physiography--of the sedimentary (unglaciated) Northern Great Plains. These views along with several similar images presented below showed the "lay of the land" of the famed "Big Sky Country" as Montana is affectionately (and rightly) known. It is generally accepted that the designation of "Big Sky Country" is traceable to the western novel, The Big Sky, written by Guthrie (1947) and subsequent novels in the "Big Sky Series" all by the same author. The Great Plains is the part of Montana most descriptively associated with the "Big Sky Country". To rangemen in general and prairiemen or grasslanders in particular there is no more aesthetically exciting and spiritually moving landscape than plains grasslands extending almost as limitlessly as the endless sky itself. Simply matchless.

Garfield County, Montana. Mid-June, late vernal aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-57 (Grama-Needlegrass-Wheatgrass). SRM 607 (Wheatgrass-Needlegrass). Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40). Silty range site, 10-14 precipitation zone, Eastern Sedimentary Plains (Ross and Hunter, 1976, p. 11). Northwestern Great Plains-Montana Central Grasslands Ecoregion 43n (Woods et al., 2002).

140. Fair details- Composition and structure of a degraded wheatgrass-needlegrass prairie with considerable decline in absolute and relative cover and general abundance of climax dominants of western wheatgrass, bluebunch wheatgrass, and needle-and-thread and commensurate increase in cover, density, etc. in Sandberg's bluegrass and Wyoming big sagebrush. This "photoplot" was on the same Northern Great Plains range that was introduced in the preceding two photographs. This range was in upward range trend with range retrogression having been the result of prior (past) overgrazing and current grazing management being of proper degree of use.

There was considerable cover of threadleaf caric sedge with much regeneration of this grasslike plant species. Increasing cover of this desirable (palatable, nutritious) Carex species was an indication of on-going range improvement via secondary plant succession. (Some specimens of threadleaf sedge growing on this range were presented immediately below.)

Garfield County, Montana. Mid-June, late vernal aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-57 (Grama-Needlegrass-Wheatgrass). SRM 607 (Wheatgrass-Needlegrass). Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40). Silty range site, 10-14 precipitation zone, Eastern Sedimentary Plains (Ross and Hunter, 1976, p. 11). Northwestern Great Plains-Montana Central Grasslands Ecoregion 43n (Woods et al., 2002).

141. Threadleaf and needle spike- Threadleaf or blackroot caric sedge (Carex filifolia) presented as two cespitose plants (first slide) and textbook image of solitary spike (second slide) growing on a degraded wheatgrass-needlegrass true prairie range converted into a grazing disclimax dominated by Sandberg's bluegrass with threadleaf sedge as local associate species. Views of this degraded range-- on which these plants and shoots of threadleaf sedge presented in the next two-slide set were growing--were in the single immediately preceding photograph and the two photographs just above that (ie. last three slides preceding these two photographs). It was explained in the two preceding captions that this degraded rangeland was in upward range trend with current moderate degree of use. Local abundance of threadleaf caric sedge with ample reproduction evidenced by younger, smaller plants was one indicator of upward trend.

Although leaves of threadleaf sedge are narrow and wirely so as to make plants of this species seem small (Forest Service, 1940, p. GL5) many individual plants of this generally cespitose species achieve large to even immense size. This author has seen individual specimens on the Northern Great Plains in Montana that were considerably larger than a full-sized pick-up bed. Some large plants of threadleaf sedge growing on the Montana plains were presented above; some of these were only about half the size of a truck bed as the biggest individuals seen by this photographer often have bare centers as larger (and, presumedly, older) plants senesce and "break up" into disjunct clonal units. Impressive by any standards, especially when the forage value, grazing tolerance, and persistence under drought are considered. Threadleaf sedge is an extremely valuable range plant. (More on this aspect in subsequent captions.)

Garfield County, Montana.Mid-June; ripening-fruit phenological stage.

142. More threaded leaves and needled spikes-A good number of sexual shoots (first slide) and closer-in views of sexual shoots with terminal spikes (second slide) of threadleaf caric sedge on a depleted wheatgrass-needlegrass range (Fair range condition class) that had been degraded into a grazing disturbance (disclimax) dominated by Sandberg's bluegrass and increased cover (to noxious plant stage) of Wyoming big sagebrush.

Increasing cover and regeneration of threadleaf sedge was an indication of improving range condition (upward range trend) through processes of secondary plant succession. The value of threadleaf sedge has been indicated to students of Range Management by the inclusion of this s as one of only two Carex species on the Society for Range Management list of 200 species of the International Range Plant Identification Contest (Stubbendieck et al., 1992, ps. 326-327). Threadleaf sedge was also included in the Forest Service Range Plant Handbook, the timeless "range plant bible", (Forest Service, 1940, p. GL5). More recent good sources for this species included: Hermann (1970, ), Hurd et al. (1998, ps. 112-113), and Wilson et al. (2008, ps. 172-173).

Garfield County, Montana.Mid-June; ripening-fruit phenological stage

143. Threaded needles- Terminal spikes on sexual shoots of threadleaf caric sedge. Two views of inflorescences (now bearing achenes) on the plants seen in the immediately preceding two photographs.

The U.S. Forest Srvice (1940, ps. GL5) described threadleaf sedge (also known variously as niggerwood, hair sedge, shorthair sedge, and blackroot sedge) as "invaluable in prevention of erosion" and as being both tolerant of grazing and drought. For contestants of the International Range Plant Identification Contest, Stubbendieck et al. (1992, ps. 326-327) described threadleaf sedge as being of Excellent to Good in forage value depending on species (eg. sheep and horses to cattle, respectively) plus being "extremely valuable early spring forage" and maintaining "...high palatability throughout the growing season". In addition, long-time experience has indicated that this is one of the graminaceous species that--like buffalograss and blue grama--"cures on the vine" (= maintains good feed quality through the dormant period and winter grazing season).

Garfield County, Montana.Mid-June; ripening-fruit phenological stage.

144. Disclimax under the Big Sky- Detriorated range on the Northern Great Plains on which potential natural (climax) vegetation was grassland dominated by western wheatgrass, bluebunch wheatgrass, and neeedle-and-thread. Other important component species of the theoretical pre-Columbian plant community included Sandberg's bluegrass, thickspike wheatgrass, threadleaf caric sedge, and Wyoming big sagebrush as themost common woody species. Silver sagebrush (Artemisia cana) was also present, but only as an occasional shrub species. On this Fair condition class rangeland there had been a drastic decline in the climax dominants, the decreasers, and large increases in the associate to minor species such as Sandberg's bluegrass and Wyoming big sagebrush. These latter species were generally increasers. Some invaders such as Japanese chess and cheatgrass or downy brome were also present. Plains pricklypear was incidental being present at sub-trace proportions.

This range was by-and-large a grazing disturbance climax of Sandberg's bluegrass with enough Wyoming big sagebrush to have become a savanna (at least to be approaching the composition and physiogonomy of a grass-shrub savanna). Present grazing intensity was moderate with appropriate distribution of grazing animals so that a slow rate of range improvement (progression of range plant community to a higher successional state) via secondary plant succession was a probable outcome. Sandberg's bluegrass disclimax could be expected to be a seral stage of the past given an adequate period measurable on human time scale (probably a number of decades but not centuries or millenia). Range degradation via overgrazing had taken place in the past. Current grazing management was proper ("wise use" manageament) with proper stocking rate, season of use, distribution of use, and kind/class or range animal. This range was being grazed by beef cattle, pronghorn (Antilocapra americana), and mule deer (Odocoileus hemionus).

It was unfortunate from the standpoint of teaching (as well as that of grazing land stewardship) that this rangeland was not in Excellent range condition beecause it presented an outstanding view of Big Sky grassland at landscape scale. Physiography (general land form and topography) of the unglaciated (sedimentary) Northern Great Plains was well represented by these photographs. The descriptive moniker of "Big Sky Country", traceable to a pastoral novel by Guthrie (1947) entitled The Big Sky (and sequels of the "Big Sky Series"), remains one of the most appropriate nicknames of any state in the Union. Another--and much older--nickname that has been applied to various areas of the Western Range and that was equally appropriate to the Montana plains is "God's Country".

Garfield County, Montana. Mid-June, late vernal aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-57 (Grama-Needlegrass-Wheatgrass). SRM 607 (Wheatgrass-Needlegrass). Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40). Silty range site, 10-14 precipitation zone, Eastern Sedimentary Plains (Ross and Hunter, 1976, p. 11). Northwestern Great Plains-Montana Central Grasslands Ecoregion 43n (Woods et al., 2002).

145. Big on the northern plains- Population of Wyoming big sagebrush (Artemisia tridentata subsp. wyomingensis) on a degraded (probably Fair range condition class) grassland in the sedimentary Northern Great Plains. Climax vegetation of this range would be either a wheatgrass-needlegrass true prairie or a wheatgrass-needlegrass-blue grama mixed prairie. Wyoming big sagebrush is certainly a member of the climax range plant community, but its over-abundance (excessive cover and density compared to potentian natural vegetation) placed it into the category of a noxious range plant (brush); that is, again, at these excessive proportions of cover.

Wyoming big sagebrush quite likely dominates more acres of land in North America than any other shrub species, if not it is a close second to whatever species does hold this record. At the cover and relative species composition on this range Wyoming big sagebrush is, as just noted, brush. This native shrub species is, of course, a valuable and highly desired range plant at proportions appropriate for management goals. Wyoming big sagebrush is particularily critical to habitat of certains such as the sage grouse (Centrocercus urophasianus), the largest grouse in North America and a magnificant bird.

Long live Wyoming big sagebrush--in proper proportions of the range plant community.

Garfield County, Montana. Mid-June.

146. Now for an herbaceous composite- Shaggy fleabane (Erigeron pumilus) on a wheatgrass-needlegrass form of true praire. There are many Erigeron species across North America, most of these being grassland species. Shaggy fleabane is a widely distributed species with a biological range from British Columbia south to the Sonoran Desert in Mexico and eastward to the grasslands of northwestern Kansas. The specimen shown here (in both side and topdown views) was growing on a range in Good condition class where it was one of several forbs from other composites to lesser spikemoss, a pteridophyte, and even the common meadow mushroom.

Garfield County, Montana. Mid-June; peak bloom stage.

147. Ending with one in Poor condition- A Sandberg's bluegrass grazing disclimax. in the Northern Great Plains. The potential naturaal (climaax) vegetation was a wheatgrass-needlegrass dominance type for which dominant decreasers were western wheatgrass, bluebunch wheatgrass, and needle-and-thread. This range had been and, obviously, was still being overgrazed so that it was a Sandberg's bluegrass-Wyoming big sagebrush grazing disclimax. About the only remnants of decreaser grasses remaining on this disturbance climax were local closerly cropped spots of the rhizomatous western wheatgrass. There was almost no threadleaf sedge left.

There were almost no forbs except the conspicuous--and highly poisonous--white locoweed (Oxytropis sericea). Details of this species' tocicity was explained in succeeding slides, but for now it was enough to give the partial quote often attributed to John Bradford, "There but by the grace of God...". Given the longterm overgrazing and current season's continuing overuse of this range one could only conclude that white or silver locoweed was not palatable to the Angus cattle grazing this particular abused range. There was paucity enough of feed that if white locoweed had any palatability at all it would have been eaten by the cattle forced to graze this horribly degraded gazing land. "There but by the grace of God..."

Garfield County, Montana. Mid-June, late vernal aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-57 (Grama-Needlegrass-Wheatgrass). SRM 607 (Wheatgrass-Needlegrass). Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40). Silty range site, 10-14 precipitation zone, Eastern Sedimentary Plains (Ross and Hunter, 1976, p. 11). Northwestern Great Plains-Montana Central Grasslands Ecoregion 43n (Woods et al., 2002).

148. Poor and poisonous- A more detailed of Northern Great Plains prairie range overgrazed to Poor range condition class with herbaceous species reduced to a disturbance climax of Sandberg's bluegrass, Japanese chess, downy brome or cheatgrass, and Wyoming big sagebrush. Potential natural vegetation of this grassland was a the wheatgrass-needlegrass rangeland cover type (Shiflet, 1994) on which western wheatgrass, bluebunch wheatgrass, and needle-and-thread were climax dominants. Sandberg's bluegrass

The showy forbs with large white infloresences was white or silver locoweed, a highly toxic poisonous forb (papilionaceous legume). White locoweed was featured immediately below. It was noted in the preceding photocaption that white locoweed was obviously not palatable to the Angus cattle forced to overgraze this abused range.

This grassland was interpreted by the author as true prairie because it did not have a major shortgrass component. This was explained in detail above in introducing the Northern Great Plains portion of true prairie. The fact that this range had been overgrazed to a shortgrass disclimax dominated by Sandberg's bluegrass (and with an increased woody component of Wyoming big sagebrush) did not change the fact that the climax range vegetation was a western wheatgrass--bluebunch wheatgrass--needle-and-thread true prairie.

Locagtion note: photographs of silver or white locoweed (Oxytropis sericea) growing on this deteriorated range were presented above (in this same section) in order that several Astragalus and Oxytropis species could be viewed together.

Garfield County, Montana. Mid-June, late vernal aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-57 (Grama-Needlegrass-Wheatgrass). SRM 607 (Wheatgrass-Needlegrass). Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40). Silty range site, 10-14 precipitation zone, Eastern Sedimentary Plains (Ross and Hunter, 1976, p. 11). Northwestern Great Plains-Montana Central Grasslands Ecoregion 43n (Woods et al., 2002).

149. Green with envy- Landscape view of a true prairie of the wheatgrass-needlegrass dominance type with a consociation of green needlegrass (Stipa viridula). Western wheatgrass was the associate species. Buffalograss was present on small, overgrazed patches that resulted from past spotgrazing. The few forb species present were scarlet globemallow (Sphaeralacea coccinea), western yarrow (Achillea lanulosa= A. millefolium subsp. lanulosa), prairie spiderwort (Tradescantia occidentalis), and hoary vervain (Verbena stricta). There were some plants of other native grass species present such as little bluestem and blue grma, but these were present at trace amounts (at most) and did not constitute anything approaching a component. Similarily, the naturalized, Eurasian, annual grass Japanese chess or Japanese brome (Bromus japonicus) was present in trace amounts so as to merit acknowlegement as a cosmopolitian invader of this region. Smooth brome and Kentucky bluegrass, the two major domestic (and widely naturalized across much of this region), agronomic forage grasses were not present in proportions (relative cover) enough to be more than incidental species of this climax grassland vegetation.

This grassland consisted of one grass layer--except where local overgrazing resulted in an infrequent shortgrass layer-- and two forb layesr. There was an irregular or sporatic, low or short forb layer comprised of scarlet globemallow. The other forb layer and the grass layer were of the same general intermediate height at plant maturity. For its simplicity of structure and botanical composition this was one of the most breath-taking grasslands ever viewed by this author. The panaramatic sweep of the dissected plains contributed to the visual grandeur of this true prairie range that was being grazed by cows and calves.

A unique combination of meterological factors (early northward shift in jet stream, wet winter, rapid runoff of snowmelt, wet spring) came together this year to produce the greatest yield of grass biomass in human memory over this portion of the Northern Great Plains. Your author and his Nikon recorded the phenomenal production and the rare beauty of this Excellent condition true prairie range.

Jones County, South Dakota. Mid-June; late vernal aspect, green needlegrass at peak standing crop and soft-dough grain stage. FRES No. 38 (Plains Grasslands Ecosystem). K-57 (Grama-Needlegrass-Wheatgrass). SRM 607 (Wheatgrass-Needlegrass). Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40). Combination of Loamy Terrace and Silty range sites (Soil Conservation Service, 1994). Northwestern Great Plains- Subhumid Pierre Shale Plains Ecoregion 43f (Bryce et al., Undated).

150. Too good not to show more- The author could not resist displaying two more landscape or panoramatic views of a green needlegrass-western wheatgrass (dominant-associate species) true prairie range. This was, "for all intents and purposes", a consociation of green needlegrass with one grass layer of midgrass height and two forb layers one of which was primarily of the low-growing scarlet globemallow, a forb more characteristic of the "shortgrass country". Other forbs--all of sparse cover--included hoary vervain, western yarrow, and prairie spiderwort. with enough combined cover to constitute a sporatic forb layer (sort of). Widely scattered plants of little bluestem and blue grama relegated these native grasses to only incidental status which was in contrast to the wheatgrass-grama-needlegrass mixed prairie cover type (Shiflet, 1994; SRM 606). Japanese chess (a naturalized Eurasian annual brome) was all but absent on this Excellent condition class range. Also, nearly absent were the invasive, agronomic grasses smooth brome and Kentucky bluegrass.

An earlier-than-typical northward shift in the jet stream following on the heels of a wet winter and spring resulted in an astoundingly moist cool-growing season (and record flooding) that produced more herbage than the oldest human residents of the region could remember. (And, as a gift from the Almighty, your ever-ready photographer was there to record the record "wild grass crop" and share the beauty with viewers.)

Jones County, South Dakota. Mid-June; late vernal aspect, , green needlegrass at peak standing crop and soft-dough grain stage. FRES No. 38 (Plains Grasslands Ecosystem). K-57 (Grama-Needlegrass-Wheatgrass). SRM 607 (Wheatgrass-Needlegrass). Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40). Combination of Loamy Terrace and Silty range sites (Soil Conservation Service, 1994). Northwestern Great Plains- Subhumid Pierre Shale Plains Ecoregion 43f (Bryce et al., Undated).

151. Northern cow pasture deluxe- Climax green needlegrass-western wheatgrass true prairie range in the eastern Northern Great Plains. This was a consociation of green needlegrass with western wheatgrass as the associate species. It was explained in the two preceding captions that other native grasses (including buffalograss, in localized spotgrazed patches, and even lesser cover of little bluestem and blue grama) and the invasive, Eurasian, annual grass Japanese brome were present at trace amounts so as to be no more than incidental species. This was especially the case for the vernal society of this range plant community.

The black dots in badkground of this wide-angle landscape view were Angus cows and calves which were fortunate indeed to graze such lush forage opn this Excellent condition class grassland. It was explained above that winter and spring weather conditions had combined to produce record yields of herbage which was shown here at peak standing crop.

Jones County, South Dakota. Mid-June; late vernal aspect, , green needlegrass at peak standing crop and soft-dough grain stage. FRES No. 38 (Plains Grasslands Ecosystem). K-57 (Grama-Needlegrass-Wheatgrass). SRM 607 (Wheatgrass-Needlegrass). Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40). Combination of Loamy Terrace and Silty range sites (Soil Conservation Service, 1994). Northwestern Great Plains- Subhumid Pierre Shale Plains Ecoregion 43f (Bryce et al., Undated).

152. Mother Nature's ripening grain crop- Sward of green needlegrass-western wheatgrass (dominant-associate species) true prairie range at peak standing crop with caryopses of green needlegrass approaching grain-ripe and shedding (maturity). Western wheatgrass was in the earlier phenological stage of pre-bloom (spikes still in the boot). Plants of buffalograss, blue grama, Japanese chess were incidental species and not visible in these two "photoquadrants". Ditto on forbs including scarlet globemallow, prairie spiderwort, hoary vervain, and western yarrow.

Herbage yield and general expression of this climax grassland were at all-time (or near it) historical heighs due to one of the wettest cool-growing seasons ever measured. This made for an outstanding example of the physiogonomy and structure of this range type.

Jones County, South Dakota. Mid-June; late vernal aspect, , green needlegrass at peak standing crop and soft-dough grain stage. FRES No. 38 (Plains Grasslands Ecosystem). K-57 (Grama-Needlegrass-Wheatgrass). SRM 607 (Wheatgrass-Needlegrass). Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40). Combination of Loamy Terrace and Silty range sites (Soil Conservation Service, 1994). Northwestern Great Plains- Subhumid Pierre Shale Plains Ecoregion 43f (Bryce et al., Undated).

153. Green on Northern Great Plains- One plant (first photograph) and a panicle (second photograph) of green needlegrass (Stipa viridula) on true prairie. Green needlegrass is one of the major, often dominant, range plants of both the true prairie and, even more so, mixed prairie. Green needlegrass is one of the major consociations of mixed prairie (Clements, 1920, p. 137). Green needlegrass is a major species (with dominant, associate, etc. status varying from range site to range site) in the following rangeland cover types (Shiflet, 1994) of the Northern Great Plains: Wheatgrass-Bluestem-Needlegrass (SRM 606), Wheatgrass-Needlegrass (SRM 607), Wheatgrass-Grama-Needlegrass (SRM 608), and Wheatgrass-Grama (SRM 609). The first two of these range types is true prairie while the latter two are mixed prairie. Green needlegrass is often the principal associate even on the Western Wheatgrass Type (610) that is comprised of this single species (an extra large-scale consociation).

Green needlegrass is a native, perennial, cool-season (festucoid) grass that is strictly cespitose (tufted), a bunchgrass species. It species range extends from the Northwest Territorities east to Illinois and south to northern New Mexico and northeastern Arizona. A good source for description of green needlegrrass from a range management perspective was the Range Plant Handbook (Forest Service, 1040, p. G121). A more ready reference was Plant Facts- Green Needlegrass (Knudson, 2005). Green needlegrass is obviously a decreaser on range sites where it is a major component species of the climax range plant community (various Natural Resources Conservation Service range site deescriptions).

Although the panicles of green needlegrass are more or less contracted and generally described as "dense" (Great Plains Flora Association, 1986, ps.1230-1231) when some of the florets open (see immediately below) the panicle (at least that portion with open florets) appears to be more open as was shown here (and below). This feature combined with the fairly long and slightly twisted awns gives a"bushy" appearance to the entire plant at the sexual stage, including at the ripe-grain stage. One of the more descriptive common names of S. viridula was feather-bunchgrass (Fernald, 1950, p. 174). The awns and calluses of green needlegrass lemmas are not mechanically injurious to grazing animals.

Shoots presented in these slides were almost a yard in height having grown in a record wet winter and spring following a preceding wet year. Red coloration in background was from inflorescences of two-groove milkvetch or two-groove poisonvetch (Astragalus bisulctus) which were shown and described above in this chapter.

Crook County, Wyoming. Mid-June.

154. Green grassy jewel- Spikelets of green needlegrass at peak anthesis. The flowers of green needlegrass are interesting. Panicles can be can be entirely cleistogamous (closed florets so as to be self-fertilized), most common; entirely chasmogamous (florets are open so that anthers and stigmas are exposed for cross-pollination), or, less commonly, mixed with chasmogamous spikelets above the lower cleistogamous spikelets.For this reason, anthers of green needlegrass are quite vriable in length with those of cleistogamous florets being much shorter (Kaul et al., 2006, p. 718). Plants that have cleistogmmy also have some chasmogamous flowers. Florets seen here were quite obviously chasmogamous (in the state chasmogamy).

Crook County, Wyoming. Mid-June.

155. Diverse true prairie- Northern Great Plains (sedimentary section)grassland complex consisting of various patches having distinctive plant communities that were dominated by one to three species of midgrasses. There were two main "patch plant communities" in the range vegetation seen in these three images. Both of these main grassland communities were dominated by midgrass species. The tan- or buff-colored areas (mid- to backgrounds of photographs) were little bluestem consociations with sideoats grama as the associate species. The dark-green areas (foreground of slides) was a mixture of bluebunch wheatgrass, western wheatgrass, needle-and-thread, and Rocky Mountain or sheep fescue (Festuca saximontana= F. ovina). There were trace amounts (relative cover) of Sandberg's bluegrass throughout this grassland mosaic though mostly in the wheatgrass-dominated patches. There were trace amounts of Japanese brome, a naturalized annual invader. Overall, this grassland was a mixed wheatgrass-little bluestem-sideoats grama form of true prairie. In this general region, little bluestem attains heights of a midgrass with mature shoots generally being of the same approximate stature as sideoats grama and western wheatgrass, classic midgrass species.

The main forbs were blacksampson or purple coneflower (Echinacea angustifolia), wavyleaf thistle and white milkwort (Polygala alba). An example of the first named in prebloom stage was in the third slide. Yucca glauca was the closest thing to a shrub species on this grassland.

Ragged hills jutting into the Big Sky in distant background made up a Breaks range site.

Powder River County, Montana. Mid-June; late vernal aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-57 (Grama-Needlegrass-Wheatgrass). 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-Montana Central Grasslands Ecoregion 43n (Woods et al., 2002).

156. Little blue patch- A consociation of little bluestem within an overall mixed wheatgrass-little bluestem-sideoats grama true prairie. Sideoats grama was the associate climax species in the little bluestem consociation. There were numerous little bluestem-dominated patches similar to this one scattered throughout the overall range plant community. Other range plants in this consociational patch were "minor players" and included bluebunch wheatgrass and western wheatgrass, the next-most abundant species therein, along with trace amounts (cover) of Yucca glauca, the only woody or woody like plant; Sandberg's bluegrass; and wavyleaf thistle.

Low, sawthooth hills behind were a Breaks range site.

This range vegetation was in the unglaciated (sedimentary) section of the Northern great Plains.

Powder River County, Montana. Mid-June; late vernal aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-57 (Grama-Needlegrass-Wheatgrass). 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-Montana Central Grasslands Ecoregion 43n (Woods et al., 2002).

157. True prairie mixture- True prairie range of little bluestem (dominant), sideoats grama (major associate), and/or bluebunch wheatgrass-western wheatgrass (minor associate species) in the unglaciated Northern Great Plains. Other grasses included Rocky Mountain fescue, needle-and-thread, Sandberg's bluegrass, and Japanese brome in that general order of general abundance and relative foliar cover. Of these latter four, Rocky Mountain fescue and needle-and-thread were the only ones of managerial importance (and that only locally). Soapweed or soapweed yucca was a semi-woody or suffructicose range plant. Forbs included wavyleaf thistle, black sampson or purple coneflower, and white milkwort. Grassland vegetation in this image was a local transition zone between patches of little bluestem consociations (sideoats grama, associate species) and patches dominated by western wheatgrass,and luebunch wheatgrass (with needle-and-thread and/or Rocky Mountain fescue as associate species).

This composite prairie vegetation--what could be dubbed a "true prairie plant mix"--consisted of two major range plant communities that made up the mosaic of this climax grassland. A consociation of little bluestem with sideoats grama as the associate species was one of these two main plant community (see immediately above). This panicoid-eragrostoid grass combination made up patches of warm-season species. The other major range community was a more diverse array of species being made up of western wheatgrass and bluebunch wheatgrass as co-dominants and needle-and-thread as the associate with some little bluestem, Rocky Mountain fescue, and Sandberg's bluegrass to "round things out". This latter community (dominated overwhelmingly by festucoid grasses) comprised patches of cool-season grasses. An example of this cool-season grass community was introduced in foregrounds of the three slide-caption two sets above. Patches of these two distinct grass communities--and relatively wide transition zones among them-- formed the vegetational mosaic of this overall range plant community that best fit the rangeland cover type described as wheatgrass-bluestem-needlegrass (Shiflet, 1994).

This climax vegetation could be termed (for teaching or illustrative purposes) a "mixed true prairie" where mixed referred to varying degrees of dominance by the 1) three main subfamilies of the Gramineae (Festucoideae, Eragrostoideae, and Panicoideae) and 2 (consequently) both cool-and warm-season grasses.

Powder River County, Montana. Mid-June; late vernal aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-57 (Grama-Needlegrass-Wheatgrass). 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-Montana Central Grasslands Ecoregion 43n (Woods et al., 2002).

158. Edge of two "patch communities"- Perimeter of transitional range vegetation between 1) a little bluestem consociation (sideoats grama, assoicate species) and 2) mixed stand of western wheatgrass and bluebunch wheatgrass (needle-and-thread and Rocky Mountain fescue, associates). Sexual shoots of bluebunch wheatgrass were in immediate foreground with sideoats grama locally dominant immediately behind in a narrow strip while the background was a consociation of little bluestem (with substantial cover of sideoats grama and much less cover of the wheatgrasses and needle-and-thread).

Details of the two "patch communities" and transition vegetation separating them was given in captions above.

Powder River County, Montana. Mid-June; late vernal aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-57 (Grama-Needlegrass-Wheatgrass). 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-Montana Central Grasslands Ecoregion 43n (Woods et al., 2002).

159. Carpet of bunchgrasses- Two "photoplots" of a little bluestem-sideoats grama (dominant-associate species) grassland stand at farther and closer distances (first and second photographs, respectively). Viewers should note that the shoots of little bluestem (distinctive buff-colored, cespitose plants) were about the same height as those of sideoats grama (smaller diameter clumps with "wood-shavings"-like basal leaves). Sideoats grama was featured in foreground of the first (more-distant) slide and to the right in the second (closer distance view) slide.

Powder River County, Montana. Mid-June; late vernal aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-57 (Grama-Needlegrass-Wheatgrass). 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-Montana Central Grasslands Ecoregion 43n (Woods et al., 2002).

160. At home near edge of its range- Several different plants of bluebunch wheatgrass (Agropyron spicatum) on a wheatgrass-needlegrass (western wheatgrass and bluebunch wheatgrass as co-dominants with needle-and-thread and Rocky Mountain fescue as associates) true prairie in unglaciated Northern Great Plains. In the plains region bluebunch wheatgrass grows as far east as the western Dakotas so plants shown here from southeastern Montana were near the perimeter of their species (biological range).

These plants were unusually large and robust having had two back-to-back years of record precipitation coupled with what appeared to be light, even extrmely light-grazing (degree of use). This may partly explain the high proportion of dead (last year's) tillers, complete with unshattered spikelets on spikes, along with large current year's tillers. Bluebunch wheatgrass is not well-adapted to heavy defoliation. Heady (1950) conducted a monographic (and classic) study of height-weight relationships of bluebunch wheatgrass from which he determined that in a typical year in eastern Montana this species should have greater than a six-inch stubble height following the grazing season. Bluebunch wheatgrass was particularily vulnerable to defoliation when it was in flower (Heady, 1950).

The extreme cespitose (tufted) habit of this bunchgrass species was conspicuous in individuals seen above. Bluebunch wheatgrass is one of the more beautiful grasses of the Northern Great Plains in addition to being one of the major range forage species. Montanans chose wisely in selecting bluebunch wheatgrass as State Grass of Montana; .certainly a plant species deserving of the Big Sky Country.

Powder River County, Montana. Mid-June; preboot phenological stage.

161. A typical bunch- An extremely robust plant of bluebunch wheatgrass on an ecotone of true prairie between communities of 1) little bluestem-sideoats grama (dominant-associate) grassland and 2) western wheatgrass and bluebunch wheatgrass (co-dominants) with needle-and-thread and Rocky Mountain fescue as associate species in the Northern Great Plains. The range on which this large specimen grew had not been grazed during the current year's plant growing season. In fact degree of use of major forage species stongly suggested that grazing had been light--extremely light as matter of fact--in the preceding year. These two consective years had received record amounts of precipittion so that comparatively great sizes of plants such as the one shown here was atypical, and likely reflected influence of plant growing conditions (ie. record soil moisture content) rather than grazing factors like stocking rate. These plants were not, however, vastly larger than those produced in typical years.

Students should note that this plant included both living current year's shoots along with dead and weeathered shoots produced last year. Astute observers will also have noted the sexual tiller complete with spike from the preceding growing season (year) in lowr right of this photograph. There were parts of seven other cespitose plants of bluebunch wheatgrass in this photograph.

Powder River County, Montana. Mid-June; preboot phenological stage.

162. This year's and last's- Bleached spikes (last year's) and green, immature spikes (this year's) of bluebunch wheatgrass (Agropyron spicatum) that was growing (and reproducing sexually) on a little bluestem-creeping juniper savanna in the unglaciated Northern Great Plains. There are from two or three up to nine spikelets per spike (Great Plains Flora Association, 1986, p. 1126). Spikes shown here were well-filled with grain-bearing spikelets.

Little Missouri River National Grassland, Golden Valley County, North Dakota. Late June; pre-anthesis stage of this year's shoots.

Note on nomenclature: Bluebunch wheatgrass is another one of the major grass species long-known (throughout most of the classic literature of grassland ecology, natural pastures, forest grazing, as well as standard manual and taxonomic treatments) by the same scientific binomial--Agropyron spicatum, in this instance--that has come in for a series of major revisions and consequent name changes. Agropyron spicatum was replaced by Elytrigia spicata which, as it turned out, was relatively short-lived so as to be superceded by Elymus spicatus which in turn was summarily replaced by the current (as of this writing) binomial, Pseudoroegneria spicata in Flora of North America (Barkworth et al. (2007, ps. 279-281), the supposed ultimate authority.

Meanwhile, current (again, of this writing) floras (or manuals) continue to use one or the other of the newer--though not necessrily the newest--scientific name. For example, the Jepson Manual- Vascular Plants of California (Baldwin et al, 2012) still showed Elymus spicatus whereas Shaw (2008) in Grasses of Colorado and Skinner (2010) in A Field Guide to Wyoming Grasses used Pseudoroegneria spicata.

163. A confusing associate- Rocky Mountain fescue (Festuca saximontana= F. brachyphylla subsp. saximontana=, in part and probably mistaken, F. ovina) growing on true prairie in a local ecotone between a little bluestem-sideots grama and bluebunch wheatgrass-western wheatgrassas community (shown and described above). The plants shown here in the immediately succeeding two slides were particularily robust and vigerous at the end of a record wet winter and spring that followied on the heels of record precipitation the preceding year. Given these nusually large sizes and the morphological closeness of this species to that of Idaho fescue (F. idahoensis) confusion between these two frequently co-existant Festuca species.

Another point of confusion related to the scientific name of Rocky Mountain fescue. Barkworth et al., (2006, p. 422) indicated that throughout much of the earlier literature (including floras and manuals) Festuca ovina was the binomial (of an introduced Eurasian species) applied to most of the finner- (narrower-) leafed Festuca species other than F. idahoensis. Some earlier authors like Rykberg (1922, p. 86) and, later, Fernald (1950, ps. 104-105) recognized and distinguished between F. saximontana and F. brachyphylla. Great Plains flora Association (1986, p. 1181) recognized F. saximontana as F. ovina var. rydbergii. Barkworth et al. (2006, ps. 428-432) provided the most detainled discussion and distinctions between F. saximontana and F. brachyphylla. Skinner (2010, p. 246-249), the grass field guide closest to location where plants seen here were photographed, also distinguished between F. saximontana and F. brachyphylla.

Powder River County, Montana. Mid-June; immediate post-anthesis stage of phenology.

164. Not sheepish- Two strongly cespitose plants of Rocky Mountain fescue on an ecotone between two true prairie communities: 1) little bluestem-sideoats gaama and 2) bluebunch wheatgrass-western wheatgrassas in the sedimentary Northern Great Plains. The first of these two photographs was taken under an overcast sky (one large cloud between sun and subject) while the second image of the same two plants was taken a few moments later when the could dissipated. Viewers should note that the color of panicles was more pronounced under full sunlight (second slide) than under overcast (first slide).

It was remarked in the preceding caption that these plants which had a height of 18 to 20 inches had grown in the second record wet spring in a row.

Powder River County, Montana. Mid-June; immediate post-anthesis stage of phenology.

165. Distinctive panicles- Panicles with general details of spkikelets of Rocky Mountain fescue produced on true prairie in the unglaciated Northern Great Plains. These were panicles (after having been freshly pressed and dried) off of plants shown in the two preceding two slide-caption sets.

Powder River County, Montana. Mid-June; immediate post-anthesis stage.

166. Related species for point of reference- Highway right of way seeding (first slide) and panicle (second slide) of Idaho fescue (Festuca idahoensis) shown for purposes of comparison to examples of the naturally occurring plants of Rocky Mountain fescue shown immediately above. These two taxa are almost indistinguishable with statue (culm and panicle length) being the main distinguishing features according to Dorn (1984, p. 187) in Vascular Plants of Montana, the only flora for the Big Sky Country as of the writing of this caption.

It was explained in a preceding caption for Rocky Mountain fescue that the species name of Festuca ovina has most likely been misapplied to what should be regarded as F. saximontana= F. brachyphylla subsp. saximontana, but for purposes of comparison F. ovina, as used in Dorn (1984, p. 187), was acknowldeged to avoid further confusion. Anyway, an example of F. idahoensis in an adjoining area was shown here for comparison to the nearly identical species known variously as Rocky Mountain and, more traditionally or historically, as or sheep fescue.

Powder River County, Montana. Mid-June; immediate post-anthesis stage.

167. Transition true prairie (one that does fit not the vegetation map)- Natural grassland co-dominated by little bluestem and needle-and-thread in a transition zone between the Nebraska Sandhills to the south and the Pierre Shale Plains and White River Badlands to the north. Junegrass as the main associate species in this late vernal-early estival society. Threadleaf caric sedge was locally common. The main forb in this seasonal aspect was silverleaf scurfpea or silverleaf Indian breadroot (Psoralea argophylla). White milkwort was an occasional forb (example included in a photograph shown below).

At large (semi-regional) scale Kuchler (1966) mapped this unit of potential natural vegetation as wheatgrass-needlegrass with dominants being western wheatgrass, blue grama, needle-and-thread, and green needlegrass (Kuchler, 1964, p. 66). On the grassland shown here there was almost no western wheatgrass (and relatively little blue grama) and, instead a co-dominant in little bluestem which was was not listed for the wheatgrass-needlegrass unit (Kuchler (1964, p. 66) and , instead, was designated as a dominant of Nebraska Sandhills prairie (Kuchler, 1964, p. 75). Needle-and-thread was listed as a dominant for both of these units which showied the affinity and unity of these two units which were described by Barker and Whitman (1989) and adopted as rangeland cover types (SRM 602 and SRM 607) in Shiflet (1994),

Similarly, the wheatgrass-bluestem-needlegrass unit of potential natural vegetation (Kuchler, 1964, p. 67) was mapped (Kuchler, 1966) to the east of--though contiguous with--the grassland vegetation presented here. This unit of natural vegetation was also described by Barker and Whitman (1989) and adopted as a rangeland cover type (SRM 606; Shiflet, 1994) It seemed apparent to the current author that the little bluestem--needle-and-thread prairie treated here was an ecotonal grassland. In this vein, it was emphasized that the highly rhizomatous western wheatgrass is generally more tolerant of heavy grazing than is the cespitose little bluestem such that it was highly unlikely that heavy grazing had eliminated western wheatgrass while sparing little bluestem. It could be postulated that heavy grazing or overgrazing during the early green feed period eliminated the cool-season western wheatgrass while leaving the warm-season little bluestem. Any such argument is illogical and fatally flawed because co-dominant needle-and-thread is also a cool-season species and one typically more susceptible to heavy defoliation than is western wheatgrass. Likewise, Junegrass is a cool-season decreaser.

Under current treatment, the author regarded this little bluestem--needle-and-thread-dominated prairie as climax grassland that fit the Clementsian distinction of true prairie in which three of the five consociation-forming, association dominants were little bluestem, needle-and-thread, and Junegrass (Clements, 1920, ps.121-128) Clements (1920, p. 121) identified true prairie as the Stipa-Koleria association.

The two photographs used to introduce this transitional true prairie showed the physiography of this part of the Northern Great Plains as well as physiogonomy and overall species composition of the range plant community.

Todd County, South Dakota. Late June-early estival aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-59 (Wheatgrass-Needlegrass). Transition of SRM 602, SRM 606, and SRM 607. Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40). Northwestern Great Plains- Keya Paha Tablelands Ecoregion 43i (Bryce et al., Undated).

168. Ecotonal true prairie- Structure and species composition of a little bluestem--needle-and-thread-dominated grassland that formed an ecotone between Nebraska Sandhills tallgrass prairie and true and mixed prairies of the White River Badlands and Pierre Shale Plains. This grassland vegetation was distinguished and defined by absence of a major western wheatgrass component (and, less diagonistically, of blue grama as a major shortgrass component) along with presence of Junegrass as associate species (at least in this spring-summer society). There was (almost always is) some blue grama, but it was clearly a minor species. Also largely absent were plants of Kentucky bluegrass and Japanese brome other than as occasional plants (ie. "few and far between"). Threadleaf caric sedge was present though abundant only at local scale (ie. a patchy distribution). The main forb was silverleaf scurfpea or slilverleaf Indian breadroot. White milkwort was another forb found infrequently (see immediately below).

Todd County, South Dakota. Late June-early estival aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-59 (Wheatgrass-Needlegrass). Transition of SRM 602, SRM 606, and SRM 607. Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40). Northwestern Great Plains- Keya Paha Tablelands Ecoregion 43i (Bryce et al., Undated).

169. Details of a simple grassland- Two "photoplots" of true prairie co-dominated by little bluestem and needle-and-thread (Junegrass was associate) in the Northern Great Plains during a subtle shift between vernal to estival aspects. The principal forb (not represented here) was silverleaf scurfpea or slilverleaf Indian breadroot. The forb featured in the second of these two slides was white milkwort. It was explained above that this true prairie range was ecotonal with tallgrass, true, and mixed prairies in this region.

Todd County, South Dakota. Late June-early estival aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-59 (Wheatgrass-Needlegrass). Transition of SRM 602, SRM 606, and SRM 607. Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40). Northwestern Great Plains- Keya Paha Tablelands Ecoregion 43i (Bryce et al., Undated).

170. Forest and grassland- A mosaic of ponderosa pine (Pinus ponderosa) pine forest with an herbaceous understorey dominated by slender wheatgrass (Agropyron trachyculum= A. subsecundum= A. caninum) and true prairie grassland of midgrasses the major species were little bluestem, western wheatgrass, needle-and-thread, and bluebunch wheatgrass (in that overall relative order). The conspicuous forb in foreground of both of these slides was large Indian breadroot, breadroot scurfpea, or prairie-turnip (Psoralea esculenta).

This range vegetation could be seen as either forest with grassland communities developing as glades within or, alternatively, as grassland with covers or groves of forest or woodland. A third perspective or mental/verbal imaging of this Northern Great Plains vegetation was that of a savanna in which groves of ponderosa pine forest comprised a woody component of the overall (at regional-scale) predominant grassland resulting in a savanna physiogonomy. Savannahs have traditionally been regarded as ecotones or transition zones between grassland and woody (tree and/or shrub) vegetation (Dyksterhuis, 1957) This composite or mosaic vegetation did not handily the accepted or classical savanna concept of Dyksterhuis(1957) because these ponderosa pine forest had developed as "vegetational islands" (isolated remnants or, perhaps, relicts of forests that formed over earlier climatic periods) and not as extensions of present era-forests.

Certainly the open canopy, herbaceous understorey, ponderosa pine forests were in monoclimax theory postclimax vegetation (a more mesic habitat-requiring plant community) within the zonal or regional present climax (climatic climax). Polyclimax or climax pattern theories could just as handily provide explanation (successional intrpretations) and description of this patchwork of range vegetation. Obviously groves of ponderosa pine forest developed on different range sites than those of midgrass grassland.

Whatever the visualizatioon or interpretation it was interesting, eductional , inspiring range vegetation. Perhaps even more to the point was the productivity of these various range plant communities and their production of landscape "goods and srvices". For example, a large flock (rafter or gang) of Merriam's wild turkey (Meleagris gallopavo merriami) were foraging at the edge of forest and grassland presented in these and subsequent slides. Unfortunately, the turkeys did not welcome intrusion by this photographer and they vamoosed before the Nikon could capture them in their habitat. Perhaps this was appropriate from the standpoint of nativeness given that Range Types of North America was devoted to native flora. It has been agreed among wildlifers that turkey was not part of the native or indigenous fauna to that part of North American that is now the state of Montana wherein this avian species was introduced by the whiteman strarting in the 1950s (Dickson, 1992, p. 374).

This mosaic of forest and grassland provided a good example of where Landscape Ecology is a more apporpriate framework then Ecosystem Ecology from which to view and analysis range (in this case both rangeland and grazable forest).

Powder River County, Montana. Mid-June; late vernal aspect. Mosaic of FRES No. 38 (Plains Grasslands Ecosystem), K-57 (Grama-Needlegrass-Wheatgrass), SRM 606 (Wheatgrass-Bluestem-Needlegrass), Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40) and FRES No. 21 (Ponderosa Pine Forest Ecosystem), K-15 (Eastern Ponderosa Forest), SAF 237 (Interior Ponderosa Pine), eastern version of SRM 110 (Ponderosa Pine-Grassland), Cold Temperate Forest and Woodland 122- Rocky Mountain Montane Conifer Forest 122.6, Yellow Pine Series 122.62 in Brown et al. (1998, p. 37). Northwestern Great Plains- Pine Scoria Hills Ecoregion 43p (Woods et al., 2002).

171. At the edge- Margins of ponderosa pine forest with understorey dominated by slender wheatgrass (background) and true prairie grassland of little bluestem, western wheatgrass, needle-and-thread, and bluebunch wheatgrass. The conspicuous and abundant forb in foreground was large Indian breadroot, breadroot scurfpea, or prairie-turnip. Microrelief resulted in a low-lying depression dominated by western wheatgrass (immediate foreground). Adjacent to and above this swale-like low area (immediate right foreground and midground) the grassland was dominated by little bluestem (buff-colored clumps) with needle-and-thread and bluebunch wheatgrass as associates. Needle-and-thread was the most widespread grass with abundant populations in both the western wheatgrass-dominated and little bluestem-dominated grassland communities.

Young ponderosa pine had invaded the grassland (covered in immediately succeeding slide-caption set).

Powder River County, Montana. Mid-June; late vernal aspect. Mosaic of FRES No. 38 (Plains Grasslands Ecosystem), K-57 (Grama-Needlegrass-Wheatgrass), SRM 606 (Wheatgrass-Bluestem-Needlegrass), Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40) and FRES No. 21 (Ponderosa Pine Forest Ecosystem), K-15 (Eastern Ponderosa Forest), SAF 237 (Interior Ponderosa Pine), eastern version of SRM 110 (Ponderosa Pine-Grassland), Cold Temperate Forest and Woodland 122- Rocky Mountain Montane Conifer Forest 122.6, Yellow Pine Series 122.62 in Brown et al. (1998, p. 37). Northwestern Great Plains- Pine Scoria Hills Ecoregion 43p (Woods et al., 2002).

172. Three zones and woody invasion- A patchwork of climax range vegetation in the unglaciated part of Northern Great Plains. In restricted areas of the northern plains a combination of true prairie grassland and groves or small stands of ponderosa pine-bunchgrass forest had developed. An example of this grassland-forest mosaic was presented in these two slides. The grassland consisted of two major communities: 1) western wheatgrass consociation with needle-and-thread as the associate speciess on low-lying local areas (depressions) and 2) little bluestem-dominated higher areas with needle-and-thread and bluebunch wheatgrass as associate species. Needle-and-thread was the most widespread grass species. In restricted local areas needle-and-thread was co-dominant with little bluestem. An example of the wesstern wheatgrass consociation was in the foreground of both slides. The little bluestem-dominated grassland was in the midground and easily recognized by the light tan- or buff-colored grass clumps. Small saplings and a few seedlings of ponderosa pine were growing on the little bluestem grassland (two paragraphs below in this same caption). Also in both grassland communities were numerous short plants of turnip-root, breadroot scurfpea, or large Indian breadroot.

The ponderosa pine-bunchgrass forest (background of both slides) had developed on the highest ground of this landscape. The forest understorey was exclusively herbaceous with slender wheatgrass as the dominant species. There were some scattered plants of the suffructicose fringed sage or fringed sagebrush in the herbaceous forest layer. The most common forb in the understorey was wavyleaf thistle.

The grassland had been (and was continuing to be) invaded by young ponderosa pines, but this was limited to the little bluestem-dominated community on higher, better-drained ground (observe carefully in both slides). Ponderosa pine typically responds best on well-drained soils (on some forest sites ponderosa pine appears tr require well-aerated, rapidly drained soils). Role of fire in maintaining this grassland and preventing development of ponderosa pine forest in this range landscape was not known to this author. It has likely received only incidental study on this series of range sites. Ponderosa pine is certainly tolerant of surface fire, but even rather low-intensity fire does thin younger pines from forests (even cool fires frequently kill a lot of smaller ponderosa pines) so as to maintain the herbaceous understorey that characterizes ponderosa pine forest across most of the biological range of this species.

Powder River County, Montana. Mid-June; late vernal aspect. Mosaic of FRES No. 38 (Plains Grasslands Ecosystem), K-57 (Grama-Needlegrass-Wheatgrass), SRM 606 (Wheatgrass-Bluestem-Needlegrass), Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40) and FRES No. 21 (Ponderosa Pine Forest Ecosystem), K-15 (Eastern Ponderosa Forest), SAF 237 (Interior Ponderosa Pine), eastern version of SRM 110 (Ponderosa Pine-Grassland), Cold Temperate Forest and Woodland 122- Rocky Mountain Montane Conifer Forest 122.6, Yellow Pine Series 122.62 in Brown et al. (1998, p. 37). Northwestern Great Plains- Pine Scoria Hills Ecoregion 43p (Woods et al., 2002).

173. Border of two range types- Outermost margin of a ponderosa pine-slender wheatgrass habitat type where it was contiguous with true prairie dominated by little bluestem and needle-and-thread and with bluebunch wheatgrass and western wheatgrass as associate species. Main forb in the grassland was prairie-turnip, large Indian breadroot or breadroot scurfpea. Details of the two plant communities that made up the true prairie grassland were described above. Detailed description of the ponderosa pine-bunchgrass grass forest followed immediately below.

Students should note the abundant crop of pine cones lying on the ground surface (right foreground) of this grassland.

Powder River County, Montana. Mid-June; late vernal aspect. Mosaic of FRES No. 38 (Plains Grasslands Ecosystem), K-57 (Grama-Needlegrass-Wheatgrass), SRM 606 (Wheatgrass-Bluestem-Needlegrass), Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40) and FRES No. 21 (Ponderosa Pine Forest Ecosystem), K-15 (Eastern Ponderosa Forest), SAF 237 (Interior Ponderosa Pine), eastern version of SRM 110 (Ponderosa Pine-Grassland), Cold Temperate Forest and Woodland 122- Rocky Mountain Montane Conifer Forest 122.6, Yellow Pine Series 122.62 in Brown et al. (1998, p. 37). Northwestern Great Plains- Pine Scoria Hills Ecoregion 43p (Woods et al., 2002).

174. Inside a forest range- Interior of a ponderosa pine-grass range in Northern Great Plains. This population of ponderosa pine was uneven-aged with all age classes represented. Larger seedlings up to mature trees were visible in these two forest "photoplots". Understorey of this range vegetation was almost exclusively herbaceous with the domi9nant being slender wheatgrass. The main forb was wavyleaf thistle. There were also a few plants of yellow sweet-clover (Melilotus officinalis), an introduced agronomic legume that has naturalized much of the Western Range, especially over most of the Central and Northern Great Plains. Fringed sage (Artemisia frigida) was a suffruticose (sub-shrub) species widespread throughout the understorey of this forest range (see below).

Powder River County, Montana. Mid-June; late vernal aspect. FRES No. 21 (Ponderosa Pine Forest Ecosystem), K-15 (Eastern Ponderosa Forest), SAF 237 (Interior Ponderosa Pine), eastern version of SRM 110 (Ponderosa Pine-Grassland), Cold Temperate Forest and Woodland 122- Rocky Mountain Montane Conifer Forest 122.6, Yellow Pine Series 122.62 in Brown et al. (1998, p. 37). Northwestern Great Plains- Pine Scoria Hills Ecoregion 43p (Woods et al., 2002).

175. Understory dominant-Slender wheatgrass ((Agropyron trachycaulum= A. subsecundum= A. caninum) that dominated the herbaceous layer of a ponderosa pine-grass forest range in the Northern Great Plains. Although slender wheatgrass tends to be a pioneer species and one generally adapted to disturbance this short-lived perennial does persist into the climax as part of the understorey of various range types (Tilley et al., 2011). Slender wheatgrass is generally regarded as excellent forage to a number of wildlife and livestock species that is adapted to a number of habitats ranging from wet to relative droughty and from open forest to hillside grasslands (Stubbendieck et al. (1992, p. 205).

Taxonomic note: agrostologists (plant taxonomists in general) have argued for decades over the proper name of slender wheatgrass with first one group of authors (Hitchcock and Chase, 1950; Great Plains "Flora Association, 1986) and then three other groups of authorities (Skinner et al., 1999; Barkworth et al., 2007; Shaw, 2008) arguing for this specific epithet or that genus, each insisting with the arrogance, smugness, and self-rightousness of taxonomic Pharisees that their approach, name, etc. is the only "way, truth, and the light". With advent and ascent of cladistics this "tempest in a teapot" grew even more acrimonious. Rangemen, foresters, agronomists, and horticulturalists are botanical heathen or Gentiles to the high priests of Plant Taxonomy prompting this nonbeliever of such nonsense to offer this, "Damn the lot of them". Call it slender wheatgrass and go on.

Powder River County, Montana. Mid-June; late vernal aspect. FRES No. 21 (Ponderosa Pine Forest Ecosystem), K-15 (Eastern Ponderosa Forest), SAF 237 (Interior Ponderosa Pine), eastern version of SRM 110 (Ponderosa Pine-Grassland), Cold Temperate Forest and Woodland 122- Rocky Mountain Montane Conifer Forest 122.6, Yellow Pine Series 122.62 in Brown et al. (1998, p. 37). Northwestern Great Plains- Pine Scoria Hills Ecoregion 43p (Woods et al., 2002).

176. Representative specimens- Three different plants of slender wheatgrass in the understorey of an open ponderosa pine forest range in the unglaciated Northern Great Plains. These plants, though of a short-lived perennial species, were part of the climax forest community. Note the needles of ponderosa pine all around.

Powder River County, Montana. Mid-June; immediate pre-anthesis stage.

177. Spiikes beneath the pines- Spike inflorescences of slender wheatgrass produced in an open ponderosa pine forest range in the Northern Great Plains. These spikes which were shown at progressively closer camera (focal) distance were a representative sample of some of those present on plants like the three shown immediately above. The first image included a few of last year's spikes which had shed their spikelets/florets. The rachis of slender wheatgrass dies not disarticulate to the extent of some other members of the Triticeae or Hordeae, wheat or oat tribe.

Powder River County, Montana. Mid-June; immediate pre-anthesis stage.

178. Pine straw and propagules with lower-growing pals- Understorey of ponderosa pine-grass forest range. Ground layer with pine "straw" (shed needles) and pine cones of ponderosa pine and lower herbaceous layer of fringed sage (Artemisia frigida) and some shoots of slender wheatgrass, the dominant of the herbaceous layer of this range plant community.

By whatever name this grass was the dominant--frequently the sole member--of the herbaceous understorey of this forest range.

Powder River County, Montana. Mid-June; late vernal aspect. FRES No. 21 (Ponderosa Pine Forest Ecosystem), K-15 (Eastern Ponderosa Forest), SAF 237 (Interior Ponderosa Pine), eastern version of SRM 110 (Ponderosa Pine-Grassland), Cold Temperate Forest and Woodland 122- Rocky Mountain Montane Conifer Forest 122.6, Yellow Pine Series 122.62 in Brown et al. (1998, p. 37). Northwestern Great Plains- Pine Scoria Hills Ecoregion 43p (Woods et al., 2002).

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, Mixed Prairie.

179. A variant form of badlands savanna- In Little Missouri River Badlands a western wheatgrass-plains 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 plains 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.

Range vegetation seen here was an example of the widely distributed plains silver sagebursh/western wheatgrass habitat type (Wambolt and Frisina, 2002, p.15).

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).

180. Where it can grow big- Large specimen (shown at two camera distances and angles) of plains silver sagebrush (Artemisia cana subsp. cana) growing on floodplain (first terrace) of Little Missouri River along with western wheatgrass and prairie sandreed. These three species (western wheatgrass was dominant; silver sagebrush was associate, prairie sandreed was "second runner-up") formed a mixed grass-shrub savanna in the Little Missouri River Badlands, unglaciated Northern Great Plains.

This savannh was the plains silver sagebrush-western wheatgrass habitat type (Hansen et al., 1995; Wambolt and Frisina, 2002, p.15) within the general Northern Mixed Prairie Region, but there were areas of true prairie (also groves of green ash [Fraxinus pennsylvanica] and woodland dominated by Rocky Mountain juniper [Juniperus scroplorum] well as of mixed prairie present. This was the case in particular inside the Little Missouri Badlands.

Note on arrangement: Rocky Mountain juniper woodland was treated in the chapter, Juniper-Pinyon Woodland, and chapter, Miscellaneous Forest Types-I, respectively (both chapters under Woodlands and Forests heading of Range Types of North America).

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

181. Inside a big 'un- Silvery interior of a large silver sagebursh (the same plant shown at greater camera distance in the immediately preceding two slides). A. cana is one of many Artemisia species. It is the major sagebrush species in more eastern (and more mesic) parts of the Northern Great Plains. Silver sagebrush has the same relationship to grass-shrub savannas in the northern plains that sand sagebrush (A. filifolia) has in grass savannas in the southern and central plains-- or so it appeared to this rambling rangeman.

Some plants of silver sagebrush, such as those growing on river bottoms like the one presented here, grow to robust size. Others are substantially smaller, even being depauperate individuals. Several such plants of smaller size, stature, etc. were shown above on mixed prairie (grassland rather than savanna) along with a close-in view of leaf arrangement on a leader of a small silver sagebrush plant.

Sagebrush specialists and affectionadoes (the latter of which included this author) recognize two species of silver sagebrush (Wambolt and Frisina, 2002, p.15) one of which is the plains silver sagebrush (Artemisia cana subsp. cana) shown here.

References- in addition to Wambolt and Frisina (2002), Montana Sagebrush Guide, the timeless classic of Beetle (1960), A Study of Sagebrush- The Section Tridentatae of Artemisia, was strongly endorsed and recommended.

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