Palouse Prairie

(including channeled scablands)

The Palouse Prairie is somewhat ambiguous in its botanical and general physical-chemical-biological relations to other range, especially grassland, communities. This bunchgrass prairie or, if preferred, a steppe of cespitose grasses has been interpreted as a Great Plains grassland (Weaver and Albertson, 1956, ps.339-343) even though it is generally west of the Rocky Mountains not to mention the bulk of the Great Plains physiographic province. Weaver and Albertson (1956, ps. 340-341) cited other range ecologists like Clements and Heady who interpreted the Palouse Prairie as originally extending (in pre-Columbian or, at least, pre-plowing time) over portions of the Northern Great Plains as far east as the present-day states of North Dakota and South Dakota. Workers like Clements (1920, ps. 149-150) concluded that there was not a clear distinction between the Palouse Prairie and the mixed prairie of the Northern Great Plains, because this "bunch-grass prairie passes so gradually into the mixed prairie..." though the more easterly parts of bunch-grass prairie existed "[i]n the form of outposts".

Likewise, workers from Clements (1920, ps. 149-152) through Dodd (in Gould, 1968, p. 333) to Heady and Heady et al. (in Barbour and Major, 1995, ps. 493-494, 734, respectively) noted the floristic affinity between the Palouse Prairie and the original (virgin, pre-white man) Pacific or California Prairie. This relationship has been expecially obvious and noted with regard to the sharing of key species, notably Idaho fescue (Festuca idahoensis) which is often dominant, even forming consociations, and, to lesser degree, California oatgrass (Danthonia californica) which extend from the Northern Rockies and central Coast Range to the extreme southern Pacific Coast of California. These two major grasslands, were often interpreted as closely related associations (Dodd in Gould, 1968, ps. 333-335) or, in case of Clements (1920,. p. 149), as one association, the Bunch-Grass Prairie (Agropyron-Stipa Association).

The Palouse Prairie also has floristic affinities and similar abiotic features with rough fescue (F. scabrella) prairie to the northwest and adjoining mountain grasslands and herbaceous understories of forests, especially those of ponderosa pine (Pinus ponderosa). The closest vegetation-relatedness, however, is with the bunchgrass shrub steppe, the savanna transition between the Palouse Prairie and the Great Basin Desert of the Basin and Range physiographic province. Affilitation of Palouse Prairie with other floras would be expected given diversity of land forms over relatively small land area (as indicated by numerous soil series, ecoregions, range sites, etc.). Even though the Palouse Prairie was originally "sandwiched" among various kinds of vegetation (and plant-animal communities), this grassland is notable for a rather limited number of plant species (for instance as compared to tallgrass prairie that also adjoined forests and had various savannas).

Traditionally the Palouse Prairie has been regarded as a separate or distinct grassland range type (regardless of hierarchial rank in classification of natural vegetation). In fact, it struck the current author as interesting (and revealing, though of what was not so obvious) that Palouse Prairie usually has been (is) capitalized in contrast to mixed prairie, rough fescue prairie, true prairie, etc. Obviously Palouse is a proper noun that has been widely used for promotional purposes including the name for a synthetic breed of hog developed by the Washington Agricultural Experiment Station. Palouse apparently applied to the Palouse Indians, a Shahapitan people named for the Palouse River and Palouse Hills of what is now eastern Washington and northwestern Idaho. This does not explain the common convention of capitalizing Prairie when accompanying Palouse (eg. Weaver and Abertson, 1956, ps. 339, 340; Shelford, 1963, p. 350). (Then there were those like Holechek et al. [1998 and later editions] who did not even capitalize "palouse".) The designation of a specific prairie by capitalization was a convention used in the American Midwest and South to designate local prairies ranging in size from some of only a few thousand acres to those of hundreds of thousands, or perhaps, more than a million acres. A wll-known example of the latter is the Fort Worth Prairie which is the northern part of the greater Grand Prairie. Examples of small or local prairies, often indicated by the surname of a prominent family of settlers included, Swars Prairie and Burkhart Prairie of Newton County, Missouri. Ironically many of the prairies indicated by proper names, including the Palouse Prairie, have been or are being destroyed by conversion to farm land for field crops and/or, ultimately, urbanization.

As was the case for the vast majority of grasslands in humid to semiarid regions of North America, most of the Palouse Prairie fell before the plow share which, peaceful or otherwise, proved to be every bit as destructive as the sword. Much of this plowing was on marginal or even submarginal land that should never have been "broke out". Rates of soil loss in the Palouse Hills by extraordinary accelerated soil erosion have been some of the greatest on Earth, but this negative externality or market failure did not serve as a meaningful warning in time to save what once were some of the most graceful and aesthetically appealing of all grasslands. Now Palouse Prairie ranges exist largely as fragments (many of such small size as to have no commercial value) except on land that is obviously non-arable (too rocky, shallow, infertile, or steep) even to an always-optimistic, scissor-billed sodbuster.

Nonetheless, the Palouse Prairie is recognized as a distinctive grassland viewed either as a large general range type or as consisting of several rangeland cover (= dominance) types. Numerous of the various range plant communities comprising the Palouse Prairie were described--usually as habitat types--by Rexford Daubenmire. Interested students should study particularily those sections dealing with grassland (vs. shrub steppe savanna) in Steppe Vegetation of Washington (Daubenmire, 1968). It was appropriate that the definitive summary source on the Palouse Prairie was that of Daubenmire (in Coupland, 1992, ps. 297-312).

The name Palouse is a varied spelling of Palus, a nomadic tribe of North American Indians. The Palus people have sometimes been associated with the Nez Perce, but accounts in the Lewis and Clark journals clearly showed these to be two separate tribes. Contrary to the standard version or legend (depending on one's pespective) the breed of horse now known as Appaloosa was more likely derived from Palouse or Palus horse than as some derivative of Nez Perce (Wikipedia). While the term Palouse Prairie originally might have been affilitated or equated with the "home range" of the Palus, it was, as explained above, quickly applied by pioneer plant ecologists to a much larger geographic region based on affilitations of plant species.

As treated herein the Palouse Prairie was interpreted so as to include the channeled scablands, a geologic land form that is adjacent to the Palouse Hills (the Palouse Prairie in its most restricted, narrow usage). Channeled scablands range was covered below under a subheading of Palouse Prairie. This organization was deemed consistent with the unique land form and geologic phenomena responsible for formation of the channeled scablands.

Palouse Prairie has sometimes been applied in a much more restrictive--and non-traditional--usage to steppe or bunchgrass prairie that is limited or close to the Palouse Hills area in the Northern Rocky Mountains in southeastern Washington and adjoining Idaho and Oregon. In this restricted meaning Palouse Prairie applies only (or almost only) to arable--even if highly erodible--land. This restricted portion of the greater Palouse Prairie is one of the most endangered ecosystems in the United States with about only one percent of the original vegetation (= virgin sod) remaining, most having been converted to farmland by turn of the Twentieth Century (Johnson and O'Neil, 2001, p. 49).

1. The Palouse Prairie- Classic bluebunch wheatgrass (Agropyron spicatum) range with this species the major dominant. Co-dominant is Idaho fescue (Festuca idahoensis); associated species are western wheatgrass, Junegrass, Sandberg bluegrass (Poa sandbergii= P. secunda). Sward is so dense that bunchgrass growth from of grasses is not obvious. Note the alien musk thistle (Carduus nutans).National Bison Range, Lake County, Montana. July. FRES No. 36 (Mountain Grasslands Ecosystem). K-43 (Fescue-Wheatgrass). SRM 101 (Bluebunch Wheatgrass). Wheatgrass Series of or within Great Basin Shrub-Grassland biotic community of Brown et al. (1998). Northern Rockies- Flathead Valley Ecoregion, 15c (Woods et al., 2002).

2. Physiogonomy of the Palouse Prairie- This solid stand of bluebunch wheatgrass-- a relict of virgin Palouse Prairie in the heart of white wheat country-- shows the characteristic appearance of bunchgrass (vs. sod-form) prairie. Range types/sites comprised of cespitose ("tufted") species are often known by the term steppe (Russian for "lowland"). Steppe is applied to grasslands with species hat have the bunchgrass habit. Such grasslands have an "open" appearance because the grasses are widely "spaced" (dispersed), often in a uniform (= regular) distribution pattern (= dispersion), with bare patches of soil between them. This spacing and its "openness" is a function of asexual reproduction by tillers rather than by tillers plus stolons and rhizomes (or seeds in annual grasses) which produces a more closed sod or turf form of prairie. This bunchgrass steppe once occurred throughout the Columbia Basin across the Cascades into more mesic areas of the Great Basin where it first formed a savanna with big sageburush (Artemisia tridentata) and other Artemesia species along with rabbitbrush (Chrysothamnus species) (ie. the sagebrush shrub steppe) and then it petered out into the Great Basin or High Desert.Here is a relict parcel of the once vast Palous Prairie.Bluebunch Wheatgrass-Sandberg Bluegrass- Idaho Fescue (in that order) type.The rest fell victim to Charlie Russell's "Trails Plowed Under" description and the former prairie now yields white wheat (Triticum aestivum).

Whitman County, Washington. July, estival aspect. FRES No. 36 (Mountain Grasslands Ecosystem; technically it is prairie not mountain but this is the FRES Ecosystem unit). Bluebunch wheatgrass consociation form of K-43 (Fescue-Wheatgrass) or K-44 (Wheatgrass-Bluegrass). SRM 101 (Bluebunch Wheatgrass). Wheatgrass Series within Great Basin Shrub-Grassland biotic community or regional formation of Brown et al. (1998). Columbia Plateau- Palouse Hills Ecoregion, 10h (McGrath et al., 2001).

The next six slides presented the landscape of some of the drier portions of the Palouse Prairie steppe to show physiogonomy, structure, species composition, and local variation of this grassland range type. Photographs were of one of the larger relicts of bunchgrass steppe vegetation in the Columbia Plateau. This range was in the Walla Walla section (20a) of the Columbia Plateaus physiographic province (Fenneman, 1931, ps. 251-251). This rangeland was in the Loess Islands Ecosystem (10b) situated among units of the Channeled Scablands (10a) and Palouse Hills (10h) Ecoregions of the Columbia Plateau.

The photographs began with a landscape-scale view with several range sites and habitat types on the rangeland within frame of the photograph and ended with photographs that emphasized species composition and diversity (or lack thereof). This range was in (part of) the Agropyron spicatum-Poa sandbergii zonal association interpreted as a climatic climax (Franklin and Dryness, 1973, p. 223). Big sagebrush, rabbitbrush species, and, except for local microsites (small patches), Idaho fescue were absent from this range (at least for all practical purposes). This was obviously steppe and not shrub-steppe range vegetation.Overall this range was an example of the Agropyron spicatum-Poa secunda habitat type or, perhaps on shallow microsites, the Agropyron spicatum-Poa secunda habitat type, lithosolic phase described by Daubenmire (1968, ps. iii, 18-20 and 47-49, respectively). This set of slides was followed by photographs of major range plant species on this outstanding remnant of Palouse Prairie grassland.

3. Across the Palouse countryside- Landscape-scale perspective of the western portion of the hill (vs. channeled scablands) phase or form of the Palouse Prairie steppe. Range in the foreground consisted primarily of ridge tops dominated by Sandberg bluegrass with basin or Great Basin wildrye (Elymus cinereus)--larger, green herbaceous clumps--growing on small depressed microsites that held more water than on adjecent habitats. A small ingeous rock outcrop was at far left midground. Slopes in background were dominated by bluebunch wheatgrass.

Adams County, Washington. June, estival aspect. FRES No. 36 (Mountain Grasslands Ecosystem). K-44 (Wheatgrass-Bluegrass). SRM 101 (Bluebunch Wheatgrass) and SRM 302 (Bluebunch Wheatgrass-Sandberg Bluegrass). Shallow range site in immediate foreground with Loamy range site in midground through distant background (Soil Conservation Service, 1967).Wheatgrass Series of or within Great Basin Shrub-Grassland biotic community or regional formation of Brown et al. (1998). Columbia Plateau- Loess Islands Ecosystem, 10b (Environmental Protection Agency, undated).

4. Bunches of bluebunch wheatgrass- Bluebunch wheatgrass was the obvious dominant of this range steppe vegetation. Interspaces among the cespitose bluebunch wheatgrass plants was populated with cheatgrass or down bromegrass (Bromus tectorum) and,secondarily, Sandberg's bluegrass. Sides of primarily north slopes in background were also dominated by bluebunch wheatgrass with more Sandberg's bluegrass and less cheatgrass than on shallow ridge tops shown here and in the immediately preceding and succeeding slides. At local scale, some of this north slope vegetation consisted almost exclusively of bluebunch wheatgrass (see two photographs immediately below), but the range vegetation shown in this photograph was the Agropyron spicatum-Poa secunda habitat type (Daubenmire,1968, ps. iii, 18-20).

Adams County, Washington. June, estival aspect. FRES No. 36 (Mountain Grasslands Ecosystem). K-44 (Wheatgrass-Bluegrass). SRM 101 (Bluebunch Wheatgrass). Shallow range site with Loamy range site in distant background, the slope (Soil Conservation Service, 1967). Wheatgrass Series of or within Great Basin Shrub-Grassland biotic community or regional formation of Brown et al. (1998). Columbia Plateau- Loess Islands Ecosystem, 10b (Environmental Protection Agency, undated).

5. Lot of individual players but just a few parts to play- The species-limited range plant community presented on this ridge top consisted of bluebunch wheatgrass, easily the dominant species, snow wild-buckwheat (Eriogonum niveum), obviously the major shrub, Sandberg's bluegrass, and cheatgrass or downy brome. Snow wild-buckwheat was the major shrub (in fact, about the only shrub of any consequence) on this range, but its status was that of an associate species not a co-dominant. This range plant community was the Agropyron spicatum-Poa secunda habitat type described by Daubenmire (1968, ps. iii, 18-20).

Adams County, Washington. June, estival aspect (dormancy in Sandberg's bluegrass, hard-grain stage in bluebunch wheatgrass, full-bloom stage in snow wild-buckwheat). FRES No. 36 (Mountain Grasslands Ecosystem). K-44 (Wheatgrass-Bluegrass). SRM 302 (Bluebunch Wheatgrass-Sandberg Bluegrass). Foreground was Shallow range site; background (slope) with Loamy range site in distant background (Soil Conservation Service, 1967). Wheatgrass Series of or within Great Basin Shrub-Grassland biotic community or regional formation of Brown et al. (1998). Columbia Plateau- Loess Islands Ecosystem, 10b (Environmental Protection Agency, undated).

6. Boring to anyone but rangeman- Consociation of bluebunch wheatgrass. The natural single-species stand was more of a population than a community from perspective of biodiversity described as to species composition. This range vegetation was a good example of Clements' consociation, a term and ecological view that accomodated the widespread phenomenon in which climax vegetation frequently consist of mostly one species. This slope was predominately north-facing and the relatively greater soil moisture content favored the larger-growing bluebunch wheatgrass over its native associate species, Sandberg's bluegrass. Cheatgrass was essentially the only other species present, and, of course, it is an alien species and not part of the natural or pre-Columbian climax plant community. Most rangemen--certainly the one who took this photgraph--could recognize Bromus tectorum as a permanent (even if new) member of the climax plant community on such Palouse Prairie grassland. Interestingly this range vegetation did not fit either either the Agropyron spicatum-Poa secunda habitat type nor the the Agropyron spicatum-Festuca idahoensis habitat type described by Daubenmire (1968).

Adams County, Washington. June, estival aspect (hard-grain stage in bluebunch wheatgrass). FRES No. 36 (Mountain Grasslands Ecosystem). K-44 (Wheatgrass-Bluegrass). SRM 101 (Bluebunch Wheatgrass). Loamy range site (Soil Conservation Service, 1967). Wheatgrass Series of or within Great Basin Shrub-Grassland biotic community or regional formation of Brown et al. (1998). Columbia Plateau- Loess Islands Ecosystem, 10b (Environmental Protection Agency, undated).

7. Lay of the land with loess- This bluebunch wheatgrass-dominated semiarid steppe provided a good example of Palouse Prairie in the Loess Islands Ecoregion (10h), complete with a coulee the bottom of which was populated by a dense stand of basin wildrye. The Soil Science Society of America (2001) defined loess as "material transported and deposited by wind and consisting of predominately silt-sized particles". Troeh et al. (1991, ps. 59-60) explained that loess deposits comprise the most widespread form of wind or aeolian depositions due to physical properties that permit silt particles to be more readily detached than clay and more easily transported than sand. Loess soils are less susceptible to wind erosion because they are low in sand content which would otherwise readily detach and break off silt particles. Conversely, loess is very prone to erosion by both water and gravity, hence both mass moverment and gully erosion.

Gullies in loess deposits and loess-based soils typically have a U-shape in contrast to most other gullies that have more of a V-shape (Troeh et al., 1991, p. 60). In the Palouse Prairie, gullies in loess soils often support dense stands of basin wildrye in their wide, flat bottoms. That phenomenon was evident in background of this photograph. Coulee is a term sometimes used in reference to deeper gullies, ravines, or draws (especially those that run water at least empemerally and have revegetated and are no longer actively eroding).

Palouse Prairie vegetation featured in the foreground of this photograph was another consociation of bluegunch wheatgrass except that Idaho fescue and Sandberg's bluegrass were somewhat more abundant than in the Agropyron spicatum consociation described immediately above. This did not fit either the Agropyron spicatum-Poa secunda habitat type nor the the Agropyron spicatum-Festuca idahoensis habitat type of Daubenmire (1968). Cheatgrass was ever present, but more as sporadic plants or local stands not a major component as on disturbed environments. About the only forb present (and it was scarce) was western yarrow (Achillea millefolium ssp. lanulosa). This was a predominately west-facing slope, but with different and deeper soil than on the north-slope rangeland shown in the preceding photograph. Range vegetation presented here was more of a mixed species stand with less domination by bluebunch wheatgrass.

Aams County, Washington. June, estival aspect (dormancy in Sandberg's bluegrass; hard-grain stage in bluebunch wheatgrass). FRES No. 36 (Mountain Grasslands Ecosystem). K-44 (Wheatgrass-Bluegrass). SRM 101 (Bluebunch Wheatgrass), overall. Loamy range site (Soil Conservation Service, 1967). Wheatgrass Series of or within Great Basin Shrub-Grassland biotic community or regional formation of Brown et al. (1998). Columbia Plateau- Loess Islands Ecosystem, 10b (Environmental Protection Agency, undated).

8. More players yet- At this location Palouse Prairie more forb species were present in the range plant community. These included narrowleaf skeleton weed or bush wire-lettuce (Stephanomeria tenuifolia var. tenuifolia), western yarrow silky lupine (Lupina sericeus), and the alien (Eurasia), naturalized weed, western or yellow salsify or goatsbeard (Tragopogon dubius). Bluebunch wheatgrass was still the dominant range plant followed by Sandberg's bluegrass, and then (and much less) cheatgrass with some Idaho fescue. This range vegetation fit the description of the Agropyron spicatum-Poa secunda habitat type (Daubenmire,1968, ps. iii, 18-20).

Adams County, Washington. June, estival aspect (dormancy to pre-bloom in the various species). FRES No. 36 (Mountain Grasslands Ecosystem). K-44 (Wheatgrass-Bluegrass). SRM 302 (Bluebunch Wheatgrass-Sandberg Bluegrass). Loamy range site (Soil Conservation Service, 1967). Wheatgrass Series of or within Great Basin Shrub-Grassland biotic community or regional formation of Brown et al. (1998). Columbia Plateau- Loess Islands Ecosystem, 10b (Environmental Protection Agency, undated).

9. A rank but spindly composite- Narrowleaf skeleton weed or bush wire-lettuce (Stephanomeria tenuifolia var. tenuifolia) grew in association with bluebunch wheatgrass on the Palouse Prairie range featured above. Adams County, Washington. June, estival aspect (pre-bloom in wire-lettuce).

10. Purty bare alright- Shoots of two plants of narrowleaf skeleton weed or bush wire-lettuce with sparse (and narrow) leaves on green shoots that were clearly capable of conducting photosynthesis. This rank-growing forb was obviously palatable as seen by the heavy degree of use on plants growing on another bluebunch wheatgrass-Sandberg's bluegrass range that had been grazed by cattle. This other range was on shallower soil and of more xeric range sites. It was shown below. The only grazing by vertebrates on the range featured here was by mule deer and rodents and rabbits. The author found no evidence of grazing on wire-lettuce plants that were growing on this range.

Adams County, Washington. June, estival aspect (pre-bloom in wire-lettuce).

11. Went all to stem- Close-up photograph of upper portion of shoots of narrowleaf skeleton weed or bush wire-lettuce. What leaves were present were quite narrow. This plant was growing on the same range as shown above. This native perennial species is in the lettuce tribe (Lactaceae) oc Compositae. Pre-bloom phenological stage. Adams County, Washington. June.

12. Western or yellow salsify or, sometimes, goatsbeard (Tragopogon dubius)- This is a naturalized, annual Eurasian weed that is common throughout much of the Palouse Prairie steppe. In fact, this member of the chicory tribe (Chicorieae) of Compositae is one of the more widely distributed exotic weeds in North America. It is not a mjor plant species on Palouse Prairie ranges and, in fact, would appear to be noteworthy only because of its prominent though infrequent appearance on this range type. It was included here as an example of another naturalized range plant and because it commonly prompts inquiry from producers, wildflower fans, and other curious parties.

Latah County, Idaho. June: phenololgy of these two plants growing on the same range varied from pre- to peak-bloom and seed-shatter.

13. Another run at it- A more robust and sexually reproductive specimen of goatsbeard (goat's beard) presented for a more "photogenic" if not better detailed view. This plant was growing by a railroad track in the west Cross Timbers of northcentral Texas. The distant separation of these two plants gave some idea of the geographic range of this naturalized composite in North America. On the other hand, this was an incomplete view given that this is an Eurasian species.

Erath County, Texas. Late April, flowering to mature fruit stages.

14. Fruits and flowers along the track- More detailed view of fruit (achene) clusters and head of Tragopogon dubius. These parts were on the same plant presented in the immediately preceding slide. Erath County, Texas. Late April, flowering to mature fruit stages.

15. Bluebunch wheatgrass (Agropyron spicatum)- Two plants of the dominant perennial bunchgrass over much of the Palouse Prairie. Bluebunch wheatgrass and Idaho fescue are co-dominants over vast areas of the Palouse Prairie and adjoining areas. They often side-by-side co-dominants, but on some range sites one or the other of these cool-season natives will form nearly exclusive consociations as if a natural monoculture or single-species stand. Idaho fescue is the more mesic and arguably the more typical dominant of the classic Palouse Prairie. Bluebunch wheatgrass appears to be dominant or associate species over a slightly greater geographic area. The cespitose nature of this species was conspicuous in these two examples. Bluebunch wheatgrass shoots are always tillers never stolons or rhizomes. These two plants were growing on the same range in the Loess Islands area described above.

Adams County, Washington. June; hard-dough to mature grain stage.

16. Snow wild-buckwheat (Eriogonum niveum)- Nice specimen of this species growing on the Loess Islands range featured above. This was the major (about the only) shrub on this form or variant of the Agropyron spicatum-Poa sandbergii zonal association. There were numerous immature individuals of this shrub speceis on this range as well as many showy (full-bloom) mature plants . Snow wild-buckwheat was locally the associate plant species, but it usually shared this distinction along with Sandburg's bluegrass. It was specified above that this climax range plant community was the Agropyron spicatum-Poa secunda habitat type not the Eriogonum niveum-Poa secunda habitat type.

Adams County, Washington. June; full-bloom phenological stage.

17. Snow wild-buckwheat- Two photographs to present details of this common dominant shrub on bluebunch wheatgrass-Sandburg's bluegrass range in the Palouse Prairie. The "peppermint candy stripe" on petals of this plant are a "dead-giveaway" as to its species. Adams County, Washington. June; full-bloom phenological stage.

18. Rangeman's Holy Grail and Holy Ground- Palouse Prairie when the Palouse had it to themselves. Relict of Pacific Interior bunchgrass steppe co-dominated by Idaho fescue and bluebunch wheatgrass but with Idaho fescue the dominant dominant (number one dominant). Sandberg's bluegrass was the associate species. Main forb was western yarrow. There were fewer plants of narrowleaf skeleton weed or bush wire-lettuce. Along perimeter of this relict range vegetation (mostly edges of a nearby road) there were some individuals of naturalized intermediate wheatgrass (Agropyron intermedium).

This was the ultimate expression of Palouse Prairie steppe vegetation. Example of the Agropyron spicatum-Festuca idahoensis zonal association regarded by Franklin and Dryness (1973, ps. 211, 216-218) as one of nine climatic climaxes in the steppe and shrub-steppe of the Collubia Basin Province. To have such a remnant of this climatic climax at landscape-scale was a treasure beyond price. A remarkable example of one of the most remarkable grasslands in North America. Clearly only a fraction of one percent of this original (pre-Columbian) form or variant of the Palouse Prairie steppe remains. Feast your eyes fellow rangemen.

Example of the Agropyron spicatum-Festuca idanoensis habitat type described by Daubernmire (1968. ps. iii, 21-23). In other words this species-dominance designation is both zonal association and habitat type.

This climax range plant community had the simplicity of only a few species. It was another example that biodiversity as measured by numbers of species, variation in species composition, etc. is often less not greater (fewer not more species) in climax than in seral stages.

Adams County, Washington. June (estival aspect); peak standing crop. FRES No. 36 (Mountain Grasslands Ecosystem). K-43 (Fescue-Wheatgrass). SRM 102 (Idaho Fescue). Loamy range site (Soil Conservation Service, 1967). Wheatgrass Series of or within Great Basin Shrub-Grassland biotic community or regional formation of Brown et al. (1998), this was actually closer to an Idaho Fescue Series which should obviously be added to an expanded Great Basin Shrub-Grassland biotic community. Columbia Plateau- Loess Islands Ecosystem, 10b (Environmental Protection Agency, undated).

19. Consociation of Idaho fescue- Another example of relict climax range vegetation in the Palouse Prairie. On a long, high right-of-way along a state highway this remnant was about all that remained of a once magnificant bunchgrass grassland that stretched horizon-to-horizon. This was also another example of the Clementsian consociation, a plant association (and, sometimes, a formation) characterized by a single dominant (Clements, 1916, p. 129). Some of the earliest work on consociations were those of Agropyron and Festuca in southeastern Washington and adjacent Idaho by Weaver (1917) and cited by Clements, 1920, p. 151) and Weaver and Albertson, 1956 (ps. 339-340).

Whitman County, Washington. June (estival aspect); peak standing crop. FRES No. 36 (Mountain Grasslands Ecosystem). K-43 (Fescue-Wheatgrass). SRM 102 (Idaho Fescue). Wheatgrass Series of or within Great Basin Shrub-Grassland biotic community or regional formation of Brown et al. (1998), but given the obvious widespread occurrence of Festuca idahoensis-dominated range vegetation throughout the Palouse Prairie the Great Basin Shrub-Grassland biotic community of Brown et al. (1998) obviously should be expanded to include an Idaho Fescue Series. That is definitely what the range plant community seen here was. Columbia Plateau- Palouse Hills Ecosystem, 10h (Environmental Protection Agency, undated).

20. Idaho fescue (Festuca idahoensis)- Habit of a plant of this cespitose species. Plants on the Idaho fescue consociation introduced in the two photographs immediately above. Whitman County, Washington. June (estival aspect); peak standing crop.

Technical note and warning to viewers: The "frosted", "sparkled", "dazzle-like", or "crystalized" appearance of leaves in grasses shown here and in some other portions of Range Types of North America was created in the scanning process by the unsatisfactory performance of an Epson Perfection V700 Photo color scanner. This scanner scanned at levels of performance that in this author's judgment varied from failure to excellent. Excellence in scanning typically occurred with JPEGs of leaves and flowers of forbs, shrubs, and most trees. Unfortunately, scanning performance was an abject failure--completely unsatisfactory in this photographer's experience--with JPEGs of grasses, grasslike plants, and, though less frequently, foliage of conifers. Any narrow leaves on plants scanned in the Epson Perfection V700 Photo color scannerin came out as "sparkles" or "crystals". This reduced crisp, "crystal-clear" 35mm Kodachrome slides to substandard photographic reproductions like those shown immediately above.

Epson Perfection V700 Photo color scanners are totally unsatisfactory for reproduction of photographs of whole and, often, even partial plants of grasses, rushes, and sedges as well as foliage of conifers.

This author took the opportunity to warn fellow photograhers and web authors against this inferior, slip-shod equipment. Do not make the mistake of investing in Epson apparatus. Furthermore, most officials higher than receptionists at the Epson Corporation were arrogant and provided no advice to this photographer on how to adjust of set the Epson Perfection V700 Photo color scanner for better results.Epson officials were jerks and the Epson Perfection V700 Photo color scanner is a piece of junk. Avoid this company and the Epson Perfection V700 Photo color scanner. Products and representatives of Epson Corporation are both inferior.

21. Panicle of Idaho fescue at full maturity- Appearance (including coloration) of the inflorescence of Idaho fescue when completely ripe and grain-shatter stage (or just before it in this instance). Latah County, Idaho. June (early summer).

22. Ripening panicles- Panicles of Idaho fescue (showing spikelets) approaching maturity. Hard-dough stage. Whitman County, Washington.

23. Consociation of basin wildrye (Elymus cinereus)- In low-lying areas (depressions, swales, small basins, potholes, hollows) within the Palouse Prairie grassland vegetation, especially in the channeled scablands, there range plant communities dominated by basin wildrye. These are consociations in the Clementsian model. In the context of his monoclimax theory Clements (1920, p. 151) interpreted basin wildrye (Elymus condensatus was the accepted binominial at that time) and, in fact, lowland plant communities in the bunchgrass prairie period, as subclimax. In the polyclimax theory depression communities dominated by basin wildrye and wetland plants are edaphic climaxes. Either way, such swale communities like the one shown are the potential natural range vegetation.

There were almost no other plant species besides basin wildrye in the swale in this photograph. There were incidental plants of Sandberg's bluegrass and cheatgrass. Many such swale communities include a layer of inland saltgrass so as to be interpreted as the Elymus cinereus-Distichlis stricta haabitat type (Daubenmire, 1968, iii, 50-51, Figure 27, p. 52), but the range vegetation presented here lacked the saltgrass component. In fact, most such basin wildrye-dominated depressions encountered and photographed by this author did not have D. stricta (at least to any appreciable extent). Presence of Bromus tectorum and absence of D. stricta might suggest or prompt speculation that the latter was grazed out and and replaced by the former, but it did seem plausible that the grazing tolerant, rhizomatous saltgrass would have been totally eliminated by overgrazing while the large (even huge) basin wildrye would have benefitted from overgrazing. It is commonly accepted that basin wildrye has a relatively high susceptability of to heavy grazing. In fact, over much of the Great Basin, Northern Rocky Mountains, and adjoining areas E. cinereus was almost eliminated by overgrazing and overmowing for hay. Rather, it seemed that this swale range vegetation (and numerous other such local swale communities as shown below) was a nearly single-species consociation much like that of bluebunch wheatgrass and Idaho fescue shown ablve.

Adams County, Washington. June (estival aspect); approaching peak standing crop. FRES No. 36 (Mountain Grasslands Ecosystem). K-43 (Fescue-Wheatgrass). For whatever reason the Society for Range Management (Shiflet, 1994) did not include any rangeland cover type designation for basin wildrye (apparently noone was willing to "take the plunge" and write it up). Alkali range site (Soil Conservation Service, 1967). Wheatgrass Series of or within Great Basin Shrub-Grassland biotic community or regional formation of Brown et al. (1998), but given the widespread occurrence of Elymus cinereus-dominated range plant communities throughout the Palouse Prairie the Great Basin Shrub-Grassland biotic community of Brown et al. (1998) should be expanded to include a Basin Wildrye Series. Columbia Plateau- Palouse Hills Ecosystem, 10h (Environmental Protection Agency, undated).

24. Minature marsh- On a local sites (more like a larger microsite) within the basin wildrye swale community introduced immediately above Baltic rush (Juncus balticus) had formed a consociation so dense as to exclude other vascular plant species except around edges of the rush stand. This local unit of range vegetation had developed within the overall or surrounding basin wildrye-dominated depression or pothole.

Adams County, Washington. June (estival aspect); peak standing crop of Baltic sedge. FRES No. 36 (Mountain Grasslands Ecosystem). K-43 (Fescue-Wheatgrass). No SRM rangeland cover type for rush (Juncus sp.)-dominated range vegetation. Fescue). Alkali range site (Soil Conservation Service, 1967). Wheatgrass Series of or within Great Basin Shrub-Grassland biotic community or regional formation of Brown et al. (1998). Columbia Plateau- Palouse Hills Ecosystem, 10h (Environmental Protection Agency, undated).

Note on organization: Lowland sites (swales, sinks, depressions, potholes, or by whatever other name) dominated by Elymus cinerus are more common on channeled scablands. Other examples of this potential natural range vegetation were included in the Channeled Scablands section below.

25. Bluebunch wheatgrass range- A consociation of bluebunch wheatgrass grew on this xeric west slope in the Owyhee Upland province (Franklin and Dyrness (1973, ps. 6, 34-38) of southeastern Oregon. This pristine stand that was nearing the end of its annual growth cycle represented some of the more westernly reaches of the Palouse Prairie (Weaver and Albertson, 1956, ps. 339-340). Most of the climax range vegetation in the Owyhee Upland and neighboring portions of the Columbia Plateau was sagebrush shrub-steppe (treated separately herein under Grasslands: Sagebrush Shrub-Steppe), but at higher elevations and on upland slopes Palouse Prairie bunchgrass steppe developed with bluebunch wheatgrass and Idaho fescue as dominants (or co-dominants) and with species like Thurber needlegrass (Stipa thurberiana) as associates. This relict vegetation was an example of such range plant communities.

Vale District, Bureau of Land Management, Malheur County, Oregon. June. Estival aspect on a xeric west slope. FRES No. 36 (Mountain Grasslands Ecosystem). K-43 (Fescue-Wheatgrass). SRM 101 (Bluebunch Wheatgrass). Wheatgrass Series of or within Great Basin Shrub-Grassland biotic community or regional formation of Brown et al. (1998). Bluebunch wheatgrass-Idaho fescue Palouse association of Kagan et al. (2004). Northern Basin and Range- Dissected High Lava Plateau Ecoregion, 80a (Thorson et al., 2003).

26. Sundown on a bluebunch wheatgrass stand- Interior view of the bluebunch wheatgrass consociation of Palouse Prairie presented in the preceding slide. Sun's evening rays accentuated and captured the aura of growing season's end for this pristine bunchgrass steppe. End of the diurnal and annual cycles was reminder of the plant succession terminus shown here and of the endless rhythm that produced such miraculous vegetation. It was range like this climax grassland that made the frontier ranching era one of the most colorful and enduring visions of American and Canadian history. It is rich, proud history of an era with a vision that has endured long after the sun set on that action-packed act of man and the land that was prelude to scientific Range Management.

Vale District, Bureau of Land Management, Malheur County, Oregon. June. Estival aspcect of a xeric west slope. FRES No. 36 (Mountin Grassland Ecosytem). K-43 (Fescue-Wheatgrass). SRM 101 (Bluebunch Wheatgrass). Wheatgrass Series within Great Basin Shrub-Grassland biotic community of Brown et al. (1998). Bluebunch wheatgrass-Idaho fescue Palouse association of Kagan et al. (2004). Northern Basin and Range- Dissected High Lava Plateau Ecoregion, 80a (Thorson et al., 2003).

27. Bluebunch wheatgrass-Idaho fescue range- An upland west slope in the Owyhee Upland province provided a favorable range site for this example of the Agropyron spicatum-Festuca idahoensis zonal association. Franklin and Dyrness (1973, ps. 211-212, 219-220) recognized this range community as one of nine zonal associations which can occur as climatic climaxes within the greater steppe and shrub-steppe zones of the Columbia Basin.

This bluebunch wheatgrass-Idaho fescue association of bunchgrass prairie is climax grassland and not the climax savanna, the ecotonal vegetation, designated as sagebrush shrub-steppe. For that reason-- and consistent with organization of range cover types herein according to biome-- this grassland plant community was included with the Palouse Prairie (of which it is most certainly a part) and not with the sagebrush shrub-steppe, within the larger spatial dimension of which it occurred. Franklin and Dyrness (1973, p. 212) distinguished among steppe, shrub-steppe, and meadow-steppe all three of which are part of and, thus, found in the Columbia Basin and some of which occur in adjacent parts of the High Lava Plains and Owyhee Upland provinces (Franklin and Dyrness, 1973, p. 6).

The sagebrush shrub-steppe constitutes such a major or rangeland cover types that it was treated separately (Grassland: Sagebrush Shrub-Steppe).

The characteristic physiognomy of bunchgrass prairie was portrayed in this textbook "photo-quadrant" of an "island" or "outlier" of climax Palouse Prairie on an upland (= higher-elevation) site. This patchwork-like pattern of major plant communities in which bunchgrass steppe occurs as relatively small units of range vegetation within the larger, surrounding, lower-elevation sagebrush shrub-steppe is similar to the mosaic of alpine communities that is above but within the zonal coniferous forest.

The habitat of this higher-elevation, west slope was more mesic than that of the surrounding lower-elevation sagebrush shrub-steppe so steppe grassland rather than shrub-steppe savanna developed. This climax range vegetation illustrated the cital concept of range site potential. The range plant community was simple in terms of species composition and structure. There were only two major plant species and both were co-dominant. A thrid plant species (and also a grass), Sandberg bluegrass, was the associate. Differences in height among the three major bunchgrasses could be viewed as conprising two layers of vascular plants. Forbs were absent for all practical purposes. There were a few small plants of big sagebrush, but these could not rationally be considered as forming shrub layer. Again, this was grassland not shrub steppe savanna.

Franklin and Dyrness (1973, ps. 214-216) presented tables that showed species compostion, cover, and constancy for the various associations of this area, including the bluebunch wheatgrass-Idaho fescue association. California workers (Young et al. in Barbour and Major, 1995) did not include steppe communities (nor allude to exitence of such vegetation) within their treatment of sagebrush steppe.

Vale District, Bureau of Land Management, Malheur County, Oregon. June. Estival aspect; peak standing crop. FRES No. 36 (Mountain Grassland Ecosystem). K-43 (Fescue-Wheatgrass). SRM 304 (Idaho Fescue-Bluebunch Wheatgrass). Wheatgrass Series within Great Basin Shrub-Grassland biotic community or regional formation of Brown et al. (1998). Bluebunch wheatgrass-Idaho fescue Palouse association of Kagan et al. (2004). Northern Basin and Range- Dissected High Lava Plateau Ecoregion, 80a (Thorson et al., 2003).

28. Bluebunch wheatgrass-Idaho fescue steppe- Interior (inside so-to-speak) of the Agropyron spicatum-Festuca idahoensis zonal association that is a climatic climax in the Columbia Basin and parts of the Owyhee Upland provinces (Franklin and Dyrness (1973, ps. 6, 211-212, 219-220). The cespitose morphology of these bunchgrasses was obvious as was the dispersion and spatial interval among indvidual grass plants. Sandberg bluegrass was the associate species. Forbs were inconsequential and consisted mostly of various composites. Structure of this range plant community was obviously simple, however pronounced differences in height among the three Gramineae species could be interpreted as making up two layers of vascular plants. There were a few plants of big sagebrush, but these were so rare that designation of a shrub layer did not seem justified.

This "island" of Palouse Prairie had developed in the Owyhee Upland province at an elevation higher than that of the surrounding sagebrush shrub-steppe. Most accurately this range vegetation was in the Owyhee Mountains in the Payette section of the Columbia Plateau (Fenneman, 1931, ps. 226, 246-247).

Vale District, Bureau of Land Management, Malheur County, Oregon. June. Estival aspect; peak standing crop. FRES No. 36 (Mountain Grassland Ecosystem). K-43 (Fescue-Wheatgrass). SRM 304 (Idaho Fescue-Bluebunch Wheatgrass). Wheatgrass Series within Great Basin Shrub-Grassland biotic community of Brown et al. (1998). Bluebunch wheatgrass-Idaho fescue Palouse association of Kagan et al. (2004). Northern Basin and Range- Dissected High Lava Plateau Ecoregion, 80a (Thorson et al., 2003).

29. Palouse Prairie at its successional pinnacle- An upland (= higher elevation) east slope opposite the bluebunch wheatgrass-Idaho fescue association shown in the two immediately preceding slides. Range vegetation on this more mesic east slope was also the climatic climax Agropyron spicatum-Festuca idahoensis zonal association. Physiognomy varied slightly from that of the opposite west slope steppe community, but the main difference was in species composition. Basin or giant wildrye (Elymus cinereus) was a component of the east slope range community (largest individual plants, bunchgrasses, in center foreground). In addition, Sandberg bluegrass was less common, but big sagebrush was more abundant and present as larger shrubs than in the west slope steppe community.

The aesthetic facet of this range was remarkable and warrented a passing comment. To a rangeman the totality of this land summed to grandeur. Sensuous value-- and it is much more than scenery alone-- of range is (always has been; always will be) one of the amenties and utilitarian features of this kind of land. Indeed the spiritual component is a human-affixed attribute of land of all kinds and uses. Plus, the aesthetic attribute of land-- range, in this instance-- is completely compatable with wise-use production of food, fiber, water, and other commodities and uses of natural resources.

As one takes in the splendor of range landscapes such as this, his mind-- unless his soul is dead-- harkens back to the pageant that played out (and will continue to play out) on this land. With any imagination at all there arise images of American Indians jumping buffalo (Bison bison) off of cliffs like the distant rimrock.

Such is the essence of intangibles that accompany those who live off of and with the land and work to leave it as good or better than they found it. Even Charlie Russell could not improve on this picture.

Vale District, Bureau of Land Management, Malheur County, Oregon. June. Estival aspect; peak standing crop. FRES No. 36 (Mountain Grassland Ecosystem). K-43 (Fescue-Wheatgrass). SRM 304 (Idaho Fescue-Bluebunch Wheatgrass). Wheatgrass Series within Great Basin Shrub-Grassland biotic community of Brown et al. (1998). Bluebunch wheatgrass-Idaho fescue Palouse association of Kagan et al. (2004). Northern Basin and Range- Dissected High Lava Plateau Ecoregion, 80a (Thorson et al., 2003).

30. "The Big Picture" of Palouse Prairie- A landscape perspective of the bluebunch wheatgrass-Idaho fescue cover type on the Palouse Prairie conveyed the essential features of a range ecosystem. An understanding of the ecosytem concept and its application to management of grazing land is essential to scientific preparation in Range Management, Forestry, and other natural resource fields. Of course, scientific management as such is not essential to proper management of ranges. The American Indians appear to have done a better job at that than the scientifically oriented, technologically advanced Europeans. But we cnnot turn back the clock, appealing though that fantasy is to cowhands, black powder and bow hunters, and weekend environmentalists. Largescale views such as this also serve to remind students and users of natural resources (that includes everyone) of the ecological fact that Nature operates at landscape-scale. To the professionally interested student this served as a reminder to study Landscape Ecology as well as Ecosystem Ecology. In this context landscape refers to the "spatial mosaic of several ecosystems, landforms, and plant communities across a defined area irrespective of ownership or other artificial boundaries and repeated in similar form throughout" (Helms, 1998). Obviously ecosystems can be as big as the size and scale at which one chooses to study (eg. a cow track to the biosphere), but the landscape concept of interacting ecosystems is another useful conceptual framework from which to apply principles and practices of grazing land management.

This landscape was that of the Owyhee Upland province with bunchgrass steppe as the climax plant community (potential natural vegetation). There were a few invading western juniper (Juniperus occidentalis) of young ages. These native conifers did not seem to be part of this native range vegetation but instead had invaded in absence of fire (ie.fire suppression "aided and abetted" by United States public land agencies). The matter of western juniper invasion and increase on juniper savanna and woodland and on sagebrush shrub steppe was dealt with in the section: Forest, Pinyon-Juniper Woodland.

Vale District, Bureau of Land Management, Malheur County, Oregon. June. Estival aspect. FRES No. 36 (Mountain Grassland Ecosystem). K-43 (Fescue-Wheatgrass). SRM 304 (Idaho Fescue-Bluebunch Wheatgrass). Wheatgrass Series within Great Basin Shrub-Grassland biotic community of Brown et al. (1998). Bluebunch wheatgrass-Idaho fescue Palouse association of Kagan et al. (2004). Northern Basin and Range- Dissected High Lava Plateau Ecoregion, 80a (Thorson et al., 2003).

31. Climax Palouse Prairie in the dormant season- Bluebunch wheatgrass, Idaho fescue, Sandberg bluegrass,and Junegrass, and squirrlltail bottlebrush share dominance in respective order. There are also scattered plants of big sagebrush (Artemisia tridentata). A 40 year exclosure. Clearly a typical bunchgrass habit form of several cespitose species. Vale BLM District. March. FRES No. 36 (Mountain Grasslands Ecosystem). K-43 (Fescue-Wheatgrass) and/or K-44 (Wheatgrass-Bluegrass): take your choice. SRM 302 (Bluebunch Wheatgrass-Sandberg Wheatgrass) or SRM 304 (Idaho Fescue-Bluebunch Wheatgrass, or variant thereof). Wheatgrass Series within Great Basin Shrub-Grassland biotic community of Brown et al. (1998). Northern Basin and Range- Owyhee Uplands and Canyons Ecoregion, 80f (McGrath et al., 2001).

32. Palouse Prairie in "mint condition"- This is Kuchler's "foothill prairie- Apyropron-Festuca-Stipa". This particular range is essentially a bluebunch wheatgrass consociation with scattered Sandberg bluegrass (Poa sandbergii= P. secunda). This relatively deep upland edaphic environment supports relatively little of the more mesic Idaho fescue. It is a textbook example of Daubenmire's Agropyron spicatum-Poa secunda habitat type (Daubenmire, 1970, ps. iii, 18-20; Mueggler and Stewart, 1980, ps. 21-22). This pristine bunchgrass range was being grazed by cattle and stands as a testament to outstanding stewardship and the success of "wise use" range management.

Madison County, Montana. June. FRES No. 36 (Mountain Grassland Ecosystem). K-56 (Foothills Prairie). SRM 101 (Bluebunch Wheatgrass). Closest fit in Brown et al. (1998) was Wheatgrass Series in Great Basin Shrub-Grassland biotic community.

33. Palouse Prairie (foothill prairie) vegetation in livestock exclosure- This example of bluebunch wheatgrass-Sandberg bluegrass habitat type has been protected from livestock grazing for 44 years. The exclosure was built in 1957, but is not deer- or rodent-proof. Compare this vegetation to that of the same habitat type that has been (still is being) grazed by cattle in the preceding slide. There is essentially no difference in species composition of the vegetation between the cattle range and that of the exclosure. The only apparent difference is degree of use and the greater proportion of standing dead herbage from the previous growing season in the exclosure. Renewable natural resources can be wisely used on a sustained yield basis. In the case of herbage (annual biomass), the feed resource cannot be stockpiled for long or so as to maintain high nutritive value indefinitely (ie. "use it or loose it"). Exclosures such as this one are an excellent means to insure relict vegetation which can then be used as a benchmark or point of reference to measure successful practice of Range Management and Forestry. Redbluff Ranch, Montana State University.

Madison County, Montana. June. FRES No. 36 (Mountain Grassland Ecosystem). K-56 (Foothills Prairie). Agropyron spicatum-Poa secunda (=P. sandbergii) habitat type (Daubenmire, 1970, ps. iii, 18-20). SRM 101 (Bluebunch Wheatgrass). Closest vegetation unit or biotic community in Brown et al. (1998) was Wheatgrass Series in Great Basin Shrub-Grassland biotic community or regional formation. Middle Rockies- Dry Intermontane Sagebrush Valleys Ecoregion, 17aa (Woods et al., 2002).

34. Rocky Mountain foothill prairie (Palouse Prairie)- This is an example of the Daubenmire Agropyron spicatum-Festuca idahoensis habitat type (Daubenmire, 1970, ps. iii, 21-23). It differs from the bluebunch wheatgrass-Sandberg bluegrass habitat type in the reversed relative abundance of the Poa and Festuca species. Idaho fescue is the most mesic or mesophytic of these three dominant foothill bunchgrasses (Weaver and Albertson, 1956, ps. 339, 341; Daubenmire, 1970, ps. 16, 21, 28-29; also by inference, Mueggler and Stewart, 1980, ps . 21, 31, as well as Clements, 1920, p. 150 ) and is more common or even dominant on more mesic sites such alluvial soils, higher elevations, north slopes, etc. In a somewhat atypical relationship, the more mesic species is more tolerant of grazing abuse due to its lower stature and much greater biomass in basal parts of the plant as well as its lower palatability (Mueggler and Stewart, 1980, p. 36). In the range community seen here bluebunch wheatgrass is overwhelmingly dominant, but as associate species Idaho fescue and Sandberg bluegrass are roughly equivalent. For whatever reason, Mueggler and Stewart (1980, p. 32-38) showed this as the Idaho fescue-bluebunch wheatgrass habitat type implying that the former was dominant but stating that the latter was "always present". According to the dichotomous key of Daubenmire (1970, p. iii) the mere presence of Idaho fescue distinguishes this as the bluebunch wheatgrass-Idaho fescue habitat type, but it is not as obvious as on the site seen in the next slide. Note the greener, more lush-like appearance of the bunchgrasses here when compared to that of the two previous slides. When the camera lense is moved but a few feet to the right on the same slope a slightly different plant community appears

Redbluff Ranch, Montana State University, Madison County, Montana. June. FRES No. 36 (Mountain Grassland Ecosystem). K-56 (Foothill Prairie). SRM is combination of 101 (Bluebunch Wheatgrass) and 102 (Idaho Fescue). Closest classification in Brown et al. (1998) was Wheatgrass Series within Great Basin Shrub-Grassland biotic community or regional formation. Middle Rockies- Dry Intermontane Sagebrush Valleys Ecoregion, 17aa (Woods et al., 2002).

35. Bluebunch wheatgrass- Idaho fescue Palouse Prairie- The same hillside slope as in the previous slide supports at microsite-scale the perennial forb, either silky lupine (Lupinus sericeus) or silver lupine (L. argenteus), and the shrub, rubber or gray rabbitbrush (Chrysothamnus nauseosus), both of which were absent from view in the last shot. Daubenmire (1970, p. iii) used presence of scattered rubber rabbitbrush as characteristic of this habitat type as distinct from the bluebunch wheatgrass-Sandberg bluegrass habitat type seen immediately before this habitat type. Mueggler and Stewart (1980, p. 33) noted that silky lupine could be a major member of their Festuca idahoensis/Agropyron spicatum habitat type and classified lupines as decreasers. Welsh et al. (1993, p. 438) pointed out that L. argenteus grades or phases into L. sericeus and L. caudatus so identification to the species level is not certain with this forb. Junegrass is also obvious in this community (center foreground).

These last four photographs were taken within a few miles of each other and within the same hour. Redbluff Ranch, Montana State University. Madison County, Montana. June. FRES No. 36 (Mountain Grassland Ecosystem). K-56 (Foothill Prairie). SRM 101 (Bluebunch Wheatgrass).

Nearest classification unit in Brown et al. (1998) was Wheatgrass Series within Great Basin Shrub-Grassland biotic community or regional formation. Middle Rockies- Dry Intermontane Sagebrush Valleys Ecoregion, 17aa (Woods et al., 2002).

36.Rocky Mountain foothill (= Palouse) prairie with co-dominance of Idaho fescue and bluebunch wheatgrass- More so than in the last example of the bluebunch wheatgrass-Idaho fescue habitat type, the two major species of the Palouse-northern Rocky Mountain region share "equal billing" on this pristine grassland being grazed by purebred beef cattle. Needle-and-thread (Stipa comata) and Sandberg bluegrass are the main associates. Note the presence of silver lupine as an indicator species of this habitat type. The lupine and the co-dominant bunchgrasses are, however, now joined by mountain big sagebrush (Artemesia tridentata var. vasyana). This vegetation appeared to "fit best" with the Artemesia tridentata/Agropyron spicatum habitat type as described for Montana by Mueggler and Stewart (1980, ps. 50-54). The key and description of this big sagebrush-bluebunch wheatgrass habitat type in northern Idaho and eastern Washington did not fit because criteria included absence of Idaho fescue and cover of big sagebrush as "abundant" while the big sagebrush-Idaho fescue habitat type called for absence of needle-and-thread (Daubenmire, 1970, p. iii).

This beautifully preserved and wisely used range is on the Sitz Angus Ranch, one of the earliest and most premier breeders of high-performance, top-quality beef cattle found anywhere. The quality of their range matches that of their cattle and this ranch is a sterling example of both sound grazing management as well as attention-to-detail animal husbandry.

Gallatin County, Montana. June. FRES No. 36 (Mountain Grassland Ecosystem), K-56 (Foothill Prairie). SRM is combination of 101 (Bluebunch Wheatgrass) and 102 (Idaho Fescue) or SRM 304 (Idaho Fescue-Bluebunch Wheatgrass). Closest vegetation (biotic community) unit in Brown et al. (1998) was Wheatgrass Series within Great Basin Shrub-Grassland biotic community or regional formation. Middle Rockies- Dry Intermontane Sagebrush Valleys Ecoregion, 17aa (Woods et al., 2002).

37. Needle-and-thread and bluebunch wheatgrass form of Rocky Mountain or Palouse Prairie- On this deep alluvial bench on the flood plain of the Madison River Stipa comata either dominates bluebunch wheatgrass or the two are co-dominant depending on local microsite. Both of these species largely exclude Idaho fescue, Sandberg bluegrass, and Junegrass. According to the dichotomous key of Daubenmire (9180, p. iii) bluebunch wheatgrass and needle-and-thread do not commonly co-exist (in fact they are absent one from the other), but as seen in this slide these two grasses do not always read Dr. Daubenmire's key! Mueggler and Stewart (1980, p. 33) appeared to treat this as a Stipa comata phase of their Festuca idahoensis/Agropyron spicatum habitat type. Clements (1920, p. 149-152) designated the Bunch-grass Prairie as the Agropyron-Stipa Association, but this included both the Palouse and Pacific sections (ie. the pre-Columbian California Stipa valley and foothill prairie having Mediterranean climate as well as the inland or Rocky Mountain/ Cascade grassland with continental climate but with wet winter). Clements (1920, p. 150-151) concluded that Stipa comata was a constant associate of the Agropyron-Festuca community, but "never a pure dominant" and not found in "pure communities". This distinctive range plant community illustrated that, like the more widely used range site approach, the habitat type method is "far from perfect" in identification of unique kinds of range.

Madison County, Montana. June. FRES No. 36 (Mountain Grassland Ecosystem). K-56 (Foothill Prairie). No SRM is specific: closest is variant of SRM 101 (Bluebunch Wheatgrass). Soil Conservation Service Silty range site (Ross and Hunter, 1976, p. 24). Closest biotic community (range vegetation ) unit in Brown et al. (1998) was Wheatgrass Series within Great Basin Shrub-Grassland biotic community or regional formation. Middle Rockies- Dry Intermontane Sagebrush Valleys Ecoregion, 17aa (Woods et al., 2002).

38. Idaho fescue grassland- This is the Idaho fescue phase of the Palouse Prairie (the Rocky Mountain foothill prairie portion thereof). It is a consociation of one of the two major dominant grass species of the Interior Northwest (vs. Pacific or Coastal Northwest) grasslands. This is a nearly "pure" or "solid" stand of Festuca idahoensis with bluebunch wheatgrass as a minor associate species and plains reedgrass (Calamagrostis montanesis ) as infrequent but third most abundant grass (eg. largest and darkest green clump at left foreground). This community qualifies as the Idaho fescue-bluebunch wheatgrass habitat type (Mueggler and Stewart, 1980, ps. 32-38). The Natural Vegetation of Montana Outline (Montana Natural Heritage Program, 1988) listed this as the "Festuca idahoensis Series under "Mid-grass Steppe Without Woody Plants" category. This swale is at footslope or base of a steep south slope from which runoff collects to competitively favor the more mesic Idaho fescue over bluebunch wheatgrass. The circular-shaped shrub in background is skunkbush sumac which, with the Idaho fescue, is a microhabitat-scale example of the skunkbush-Idaho fescue (Rhus trilobata/Festuca idahoensis) habitat type. Immediately upslope from this is a "regular size" mountain big sagebrush-bluebunch wheatgrass (Artemesia tridentata/Agropyron spicatum) habitat type. This latter habitat type was presented under the shrubland cover types slides.

FRES No. 36 (Mountain Grasslands Ecosystem). K-56 (Foothills Prairie). SRM 314 (Big Sagebrush-Bluebunch Wheatgrass). Closest classification or vegetation unit in Brown et al. (1998) was Wheatgrass Series in Great Basin Shrub-Grassland biotic community. Soil Conservation Service Silty range site (Hunter and Ross, 1976, p. 24). Madison County, Montana. June. Middle Rockies- Dry Intermontane Sagebrush Valleys Ecoregion, 17aa (Woods et al., 2002).

***Note to students concerning habitat types: it must be remembered that the Daubenmire habitat type— (that is the Daubenmire habitat name) and in contrast to the original habitat type of George Nichols (1917, ps. 309, 310, 317-318)— is composed of and typically bears the names of two species (Daubenmire, 1952, esp. ps. 302-303; Pfister et al., 1977, p. 9). The first species name is that of the dominant upperstorey species (in forest and shrubland habitat types this is the dominant climax tree or shrub, respectively). The second species name is that of the understory dominant species (in forest habitat types this is typically a shrub, graminoid, or, less commonly, forb species; in shrublands this is usually a grass or grasslike species). In grasslands, the dominant species for the Daubenmire habitat types is not so obvious, at least much less obvious for communities having less conspicuous layers of vegetation (eg. tallgrass prairie or shortgrass plains in contrast to mixed prairie). When one plant species forms a consociation or two species are co-dominant habitat type nomenclature becomes problematic to say the least. The example of Palouse Prairie grassland naturally dominated exclusively by one species raises questions as to either: 1) the actual meaning of habitat type (again by definition there is one dominant species in the upperstory and at least one dominant species in the understorey ) or 2) whether or not a given community or stand of essentially one species and with only scattered individuals of other species is a member of any habitat type. The examples of Idaho fescue grassland shown in the immediately preceding and the next photograph illustrate this conundrum.

39. Physiogonomy of Idaho fescue grassland- This stand (population) of Idaho fescue illustrated the outer appearance of bunchgrass vegetation (grassland composed of cespitose grasses) of the Palouse Prairie and adjacent Rocky Mountain and Cascade Range. Powell County, Montana. June.

40. Cespitose (bunched or tufted) habit- This attractive Idaho fescue plant is a textbook example of the cespitose habit of bunchgrasses. This is the origin of the term "bunchgrass prairie" for both the Palouse Prairie and the Pacific Prairie, the latter of which is dominated by Stipa species. In contrast to sod-forming grasses, bunchgrasses typically lack stolons and rhizomes (intervaginated shoots that grow horizonally) and instead have only tillers (intravaginated— thus, vertical— shoots).

***Related note and clarification: while the foothill prairie grasslands of western Montana and Wyoming are not presently conterminous with the actual Palouse Prairie region of eastern Washington and adjacent northern Idaho, historically these Festuca-Agropyron-Stipa bunchgrass prairies have been interpreted as "islands" of the Palouse Prairie. These more eastern "outposts" of the Inland and Coastal Pacific bunchgrass prairies are in effect geologically relict vegetation from when the Interior Pacific bunchgrass prairie extended as far east as the Black Hills of the Dakotas (Weaver and Albertson, 1956, ps. 339-342). Scrutiny of the Kuchler (1964, 1966 ) maps of potential natural vegetation reveals the geographic proximity and similar composition of these once extensive temperate, festucoid grasslands.

41. Sward of Idaho fescue- Two successively closer-in perspectivess showing the bunched turf of a local consociation of Idaho fescue in the Northern Rocky Mountains. This "photo-quadrant" was in a fencerow protected from livestock grazing. The cattle range immediately behind the protecting fence had been converted into a disturbance climax of the introduced agronomic pasture grass, Kentucky bluegrass (Poa pratensis).This sward was typical of the structure and physiogonomy of idaho fescue-dominated bunchgrass prairie.

Cascade County, Montana. June; ripening-grain to ripe-grain stage of phenology.

42. Ripening specimen- A relatively large plant of Idaho fescue growing in the sward portrayed immediately above. Spikelets in the hardening panicles approached grain-ripe stage of phenological development. Notice the difference in coloration of shoots in this and plants in the preceding photograph in contrast to those above and below that were in an immediate pre-bloom stage.

Cascade County, Montana. June; ripening-grain to grain-ripe phenological stage.

43. Rocky Mountain shrub steppe- This north slope beside the Madison River supports a shrub-dominated community more mesic than the benches below it but above the flood plain. The latter habitat supports grassland communities dominated variously by bluebunch wheatgrass, Idaho fescue, and needle-and-thread. Neighboring riparian vegetation varies from hydric or aquatic herbaceous species like cattails to woody plants such as willows. The shrubland or shrub steppe seen here is a transect-view going from Rocky Mountain or river juniper (Juniperus scopulorum) and big sagebrush, which appeared to be the mountain big sagebrush (Artemesia tridentata var. vasyana) form, in a breaks or rock outcrop habitat moving downslope to a mountain big sagebrush-skunkbush sumac-bluebunch wheatgrass-Idaho fescue shrub steppe. Plains reedgrass (Calamagrostis motanensis) is the main associate species of the herbaceous understory. Phenological development was slower on this north slope and at this stage bluebunch wheatgrass appeared to slightly more common than the co-dominant Idaho fescue, but this was by no means obvious grass development being far from peak standing crop. Likewise, dominance of skunkbush or big sagebrush would depend on what points along this imaginary transect were used as the basis for cover determination and species composition. Thus it would be a subjective call as to whether this was an Artemesia tridentata-Festuca idahoensis or an A. tridentata- Agropyron spicatum habitat type, or if it was a Rhus trilobata (= R. aromataca)-A. spicatum or R. trilobata-F. idahoensis habitat type. Whichever multiple guess is chosen, this is still an example of the bunchgrass shrub steppe. This raises a final question as to whether this is grassland or shrubland. Perhaps a designation of bunchgrass-shrub savanna suffices. Decisions, decisions...

FRES No. 29 (Sagebrush Ecosystem). K-49 (Sagebrush Steppe). SRM 314 (Big Sagebrush-BluebunchWheatgrass) or SRM 315 (Big Sagebrush-Idaho Fescue). Closest vegetation unit in Brown et al. (1998) was Mixed Bunchgrass-Shrub Series within Great Basin Shrub-Grassland, except Great Basin types would not apply to the Northern Rocky Mountains. Madison County, Montana. June. Middle Rockies- Dry Intermontane Sagebrush Valleys Ecoregion, 17aa (Woods et al., 2002).

44. A large relict- Large expanse of a cattle (also any wildlife such as deer and pronhorn) range that was a drier range site in the Townsend Basin on which a consociation of bluebunch wheatgrass had developed. Green needlegrass (Stipa viridula) and Idaho fescue were also present though hardly at proportions (cover, density, etc) to qualify as associate grass species. Wyoming big sagebrush (Artemisia tridentata subsp. wyomingensis) and plains pricklypear (Opuntia polycantha) added a shrub component which was present as two interrupted or sporadic shrub layers. This livestock and big game range served as an ideal or perfect example of the bunchgrass grassland of the Palouse Prairie. Range vegetation presented here and in slides immediately below was a tract of relict or virgin range vegetation even though it was being grazed by large ruminants (native and domesticated). Superb example of wise use management; outstanding range conservtion. An example of perfect stewartship of the natural grazing resources.

This relict area served as a range reference area, a range that was--from we can tell after the fact--representative of the pre-Columbian range vegetation on this part of the Northern Great Plains. Although the botanical composition or plant species diversity was low in this range plant community it was the climax vegetation and in Excellent range condition class. Species diversity along with community composition and structure are often low and more homogeneous than hetrogeneous in climax range vegetation. Students must understand the concept of the climax consociation in which one plant species is the clear dominant and, not infrequently, almost the sole species. For example Clements (1920, p. 150) specified that bluebunch wheaqtgrass "…is the major and often the exclusive dominant throughout the Palouse [Prairie], southward into Oregon and California and eastward into Idaho and Montana".

The first of these three slides was a scanned image that accurately reproduced features (colors, clarity, etc.) of plants, soil, and sky whereas the second and third slides were botched successively worse by an Epson Perfection 700 scanner. The third slide was nearly worthless, but it did covey the idea (though certainly the image) of general size and green color of a population of bluebunch wheatgrass. The third slide (which was a nearly perfect image with good depth of field before Epson Perfection "perfected" it) also included several plants of plains pricklypear in the foreground and a large plant of Wyoming big sagebrush in midground.

Although this bit of Palouse Prairie was in the Northern Rocky Mountain physiographic province (Fenneman, 1931, ps. 184, 214-215) and not the Northern Great Plains it was still part of the Palouse Prairie. Natural vegetation and physiographic provinces are not exact fits just as range vegetation and climate do not coincide or correlate perfectly. This part of the Northern Great Plains and Northern Rocky Mountains adjoin and form enclaves with each other (see Figure 82, p. 220 of Fenneman, 1931).

Townsend Basin or Valley, Lewis & Clark County, Montana. Late June; estival aspect. FRES No. 38 (Plains Grassland Ecosystem). K-59 (Wheatgrass-Needlegrass). SRM 101 (Bluebunch Wheatgrass) as applied to the Northern Rocky Mountain Cover Types. In Brown et al. (1998, p. 40) it would (there should) be Wheatgrass Series 142.1(say, 4) of Plains Grassland 142.1 of Cold Temperate Grassland 142 of Grassland Formation 140. Foothills & Mountains- Limy-Shallow-Very Shallow range site complex- 10-14 inch precipitation zone. (Ross and Hunter, 1976, ps. 27-28). Middle Rockies- Townsend Basin Ecoregion 17w (Woods et al., 2002).

45. Outward and structural appearances- Physiography, structure and overall composition of a bluebunch wheatgrass consociation of the Palouse Prairie in Excellent range condition class. Green needlegrass and Idaho fescue were secondary grass species hardly worthy of being designated as associates, but these three decreaser grasses comprised almost all of the herbaceous biomass. Cheatgrass (Bromus tectorum) was the same as non-existent on this range. This range vegetation was "el primo".

The first of these two slides showed plants of Wyoming big sagebrush, which was the major shrub (of two species for all practical purposes), and moderate degree of use around the sagebrush on this cattle range. The second slide provided an ungrazed "photoplot" (part of the range that had received very little use by cattle or wildlife) for comparison. This second slide illustrated the large-sized and rank-growing habit of which the midgrass bluebunch wheatgrass is capable. Green needlegrass, some of which were present in this view of the sward, can grow to similar size and dimensions. Idaho fescue is typically shorter in stature and generally of smaller size.

This range plant community had developed in the Northern Great Plains adjoining the Northern Rocky Mountains. Grasslands and forests develop in close proximity in this part of the Interior Northwest.

46. Internal appearances- Interior of a bluebunch wheatgrass-dominated Palouse Prairie cattle range in the Townsend Basin or Valley. The first of these two slides presented degree of use (and cattle dung showing that cattle had grazed this range) of bluebunch wheatgrass by including both representative degree of use and ungrazed bluebunch wheatgrass plants. The second slide featured mostly ungrazed plants of bluebunch wheatgrass. There was considerable range in size, both diameter of these cespitose plants as well as in height of their tillers. It was possible that larger plants were older while individuals of samller size were younger (perhaps the progeny of the larger individuals).

It was even more likely that larger plants were wolf plants that had not been grazed for as many as several consecutive years. Wolf plants in this context are those individual plants that, though of a species that is generally palatable, are not grazed by range animals (Kothman, 1974). Kothman (1074) also defined wolf plants as isolated plants that have grown to unusually large size often due to lack of or limited competition. This second meaning was less likely, but on bunchgrass prairie individual cespitose plants are often of such wide spacing that there might, in fact, be very little competition (as for water or soil nutrients for instance).

47. Internal structure- Structure and species composition of a cattle and big game range on the Palouse Prairie that was a consociation of bluebunch wheatgrass. This range plant community was almost a single-species stand of this decreaser dominant of the Interior Pacific, cool-season, bunchgrass range type. The first (horizontal) slide was a slanted downward view through this bunchgrass prairie to shown the dispersion pattern and habit of range plants. The second (vertical) slide was a top-down view showing structure, including widely-spaced dispersion, of the climax vegetation of this range, a range that was in Excellent range condition class. The second photograph also showed plants of plains pricklypear and those of the smaller Idaho fescue.

The high proportion or percentage of bare ground seen in these two slides was typical of that on northwest bunchgrass prairie. It is the nature of certain consociations, especially those with cespitose species and that are in essentially virgin or pristine condition, to have considerable area of bare soil surface and low numbers of plant species (low biodiversity; a simple species composition).

This range plant community was on the Northern Great Plains in close proximity to the Northern Rocky Mountains (Fenneman, 1931, ps. 184, 220). Such natural vegetation is equally "at home" on the plains and in the foothills of the Northern Rockies.

"I'm in love with Montana. For other states I have admiration, respect, recognition, even some affection. But with Montana it is love. And it's difficult to analyze love when you're in it."-- John Steinbeck, Travels with Charley: In Search of America

Navigational (and Ecological) note for locations in Range Types of North America: in the short section of Palouse Prairie bunchgrass steppe that follows--as well as in similar sections variously throughout this publication--there were presented grasslands that could arbitrarily (though with ecological consistency) be interpreted as either true prairie or the Stipa-Koeleria Association (Clements, 1920, 121-131), mixed prairie or Stipa -Bouteloua Association (Clements, 1920, ps. 135-139), (and/or) bunchgrass prairie or Agropyron-Stipa Association (Clements, 1920, 149-152). Or, still yet, another interpretation is that such grasslands were ecotonal (ecotonal or transitional) grasslands between any two (or, maybe, all three major Clementsian asociations.

Specifically, Clements (1920, ps. 122, 137, 150) recognized Stipa comata and Koeleria cristata consociations in the true prairie, mixed prairie, and bunchgrass prairie. Given that these two species were associates to local dominants in the example that follows various interpretations were valid. Nonetheless, the presnece of bluebunch wheatgrass (Agropyron spicatum) as overall dominant to these two species placed the following example most conssitently in the bunchgrass prairie, aone of the major forms comprising the Palouse Prairie.

"The bunchgrass prairie passes so gradually into the mixed prairie in central Montana, that no line can be drawn between them. This is readily understood when it is know that Stipa comata, Koeleria cristata, and Agropyron glaucum [smithii] occur in both, and that a large number of the societies are identical" (Clements, 1920, p. 150). Weaver and Albertson (1956, p. 340) inserted this quote in their discussion of the Palouse Prairie in Montana.

To further complicate objective interpretation, there were areas on the range presented in the following example in which Wyoming big sagebrush (Artemisia tridentata wyomingensis) was co-dominant with buebunch wheatgrass so as to form a local example of the sagebrush shrub steppe. This Wyoming big sagebrush-bluebunch wheatgrass community was probably, best regarded as a savanna form of thePalouse Prairie, the Clementsian Agropyron-Stipa Association (Clements, 1920, 149-152).

For students' convenience, the example of Palouse Prairie bluebunch wheatgrass--needle-and-thread--Junegrass steppe or bunchgrass prairie treated here was also included in the grassland chapter entitled True Prairie, given its affinity with that the Stipa-Koeleria Association (Clements, 1920, 121-131) from standpoints of physiogonomy and cover composition by the same dominant climax grass species. Given absence of Bouteloua species and buffalograss (Buchloe dactyloides) this author could not in his rangeman's mind justify it as a form of mixed prairie, the Stipa -Bouteloua Association (Clements, 1920, ps. 135-139).

48. Charlie Russell and Will James Cow Country- Landscape-scale view (first slide) and range site-scale view (second slide) of the true prairie or bunchgrass form (take your choice) of Palouse Prairie. In the concept and parlence of Landscape Ecology there is a patch of Wyoming big sagebrush-bunchgrass savanna within a matrix of true prairie in the first slide while the second slide presented a closer-in view of the true prairie matrix.

This primarily grassland range type had a bunchgrass physiogonomy and an herbaceous composition in which bluebunch wheatgrass was the overall dominant with needle-and-thread (Stipa comata) and Junegrass (Koleria cristata) as overall associates and local dominants. Indian ricegrass (Oryzopsis hymenoides) was a distant fourth-most abundant climax decreaser grass on thie pristine sample of the Palouse Prairie. There were a few widely scattered plants of threadleaf caric sedge (Carex filifolia). There were almost no forbs in this range plant community except for a few scattered plants of tufted milkvetch (Astragulus spatulatus) where native and domestic grazers could not reach them. (An example was included at end of this short section.)

This cattle range obviously was in Excellent range condition class; grassland in the same state as when Columbus' ships made landfall in this "new land for new people". This range was still native grazing land as the redmen knew it.

The rangeland shared here was a remnmant of the land that Charles Marion Russell and Will James immortalized in their artistic genuis of literature, sculpture, sketches, and paintings. It is a given that these two famed chroniclers of the frontier Big Sky Country would tip their hats to the owner of this private property just as did this photographer. Salute for flawless stewardship of these range resources.

Golden Valley County, Montana. Mid-June; vernal aspect. FRES No. 38 (Plains Grassland Ecosystem). K-59 (Wheatgrass-Needlegrass). SRM 607 (Wheatgrass-Needlegrass), bluebunch wheatgrass variant. In Brown et al. (1998, p. 40) it would (there should) be Wheatgrass Series 142.1(say, 4) of Plains Grassland 142.1 of Cold Temperate Grassland 142 of Grassland Formation 140. Western Sedimentary Plains- Silty range site complex- 10-14 inch precipitation zone (Ross and Hunter, 1976, p. 18). Northwestern Great Plains- Unglaciated Montana High Plains Ecoregion 43o (Woods et al., 2002).

49. Short ride to a rise of pure Palouse Prairie- A Palouse Prairie cattle range dominated by bluebunch wheatgrass with needle-and-thread and Junegrass as associates (locally co-dominants) and with Indian ricegrass as a fourth-ranking major grass species. Threadleaf caric sedge was present as a few scattered plants, but there were almost no forbs other than a few specimens of the nodulated legume, tufted milkvetch (shown below).

50. A savanna form- On a Palouse Prairie cattle range in the western unglaciated Northern Great Plains there were some patches of Wyoming big sagebrush in a range plant community that otherwise was a bunchgrass grassland (steppe of cespitose grasses) dominated by bluebunch wheatgrass with needle-and-thread and Junegrass as its associates. (Indian ricegrass and threadleaf caric sedge comprised most of the rest of the graminoid component of this range vegetation.) This natural vegetation was a grassland on which Wyoming big sagebrush along with perennial bunchgrasses formed a savanna rather than a big sagebrush-bunchgrass shrub steppe.The overall physiogonomy and composition of this savanna was shown in the first slide that introduced this section (two slide-caption units immediately above this slide-caption unit).

In context of Landscape Ecology the savanna form of this Palouse Prairie vegetation constituted grass-shrub patches in a surrounding grassland matrix. On closer observation it was apparent that the Wyoming big sagebrush "contingent" consisted of scattered individual plants of this shrub surrounded by the overwhelming graminoid component.

This was pristine range vegetation that was presumedly in the same succcessional status as it was before the whiteman overstocked with cattle, sheep, and horses such Big Sky ranges. Joyfully, there are still some such ranges (such as the one shared here) that are in the same ecological state as when artists like Charlie Russell recorded the romantic, idylic, and rapidly fleeting approach to livestock production in which buckeroos and sheep-herders were knights roaming freely on endless, unfenced grazing land. Also, happily, there are still a few of those vagabond, mounted cavaliers to adorn such worthy grand pastures.

51. Out of reach in safely in the rock- Beautifully blooming specimen of tufted milkvetch (Astragulus spatulatus) on a rock formation on an island of Palouse Prairie. The cattle range on which this rock outcrop and it blooming beauty occurred was dominated by bluebunch wheatgrass. This plant was growing high in its rocky refuge and beyond reach of the longest neck and most acrobatic tongue of a cow brute.

There are a great number of Astragalus species across the Western Range Region, many of which are poisonous. There is no documented evidence of stock-poisoning by A. spatulatus as shown in the encyclopedic treatment by Burrows and Tyrl (2013).

Heads of papilionaceous flowers (second or lower slide) are borne on long stalks from relatively compact plants (first or upper slide).

Golden Valley County, Montana. Mid-June; and no doubt but what this example was at peak-bloom stage.

52. Where plains and mountains meet- A Palouse Prairie range co-dominated by bluebunch wheatgrass and Idaho fescue with needle-and-thread and green needlegrass as associate species. For all practical purposes there were no forbs on this bunchgrass prairie. The one forb species found by your author was Plains phlox (Phlox andicola).

This range was in "mint condition": the climax range plant community in Excellent range condition class. The cattle were mature brood cows in late term pregnancy ("heavy springers"). A pastoral secne that restores the soul of any rangeman from King David on down. This range was soso subject to grazing by mule deer, white-tailed deer, and elk.

Park County, Montana. Late June; early estival aspect with two co-dominant grasses at peak stading crop with ripening grain. FRES No. 36 (Mountain Grasslands Ecosystem. K-43 (Fescue-Wheatgrass). SRM 304 (Idaho Fescue-Bluebunch Wheatgrass). No biotic community designation in Brown et al. (1998). Range site: Foothills and Mountains- Silty, 15-19 inch precipitation zone (Ross and Hunter,, 1976, p. 24). Northwestern Great Plains- Non-calcareous Foothill Grassland ecoregion 43s (Woods et al., 2002).

53. Overall and up closer- Superb bunchgrass steppe in part of the Palouse Prairie in the far-western Northern Great Plains at the base of the Northern Rocky Mountains (just at edge of the foothills). This climax range vegetation (Excelent range condition class) was do-cominated by Idaho fescue and bluebunch wheatgrass with green needlegrass and needle-and-thread as associate species. The author found no forbs except a few widely scattered plants of plains phlox.

Students should note the highly tufted (cespitose) habit of these bunchgrasses. This distinctive and characteristic morphological feature was also presented in succeeding slides.

54. Varied fields of view- Three photographs of climax Interior Pacific bunchgrass (steppe) prairie on a cattle range (with mule and white-tailed deer and elk being native ruminants free-ranging on this pasture) at the juncture of the Northern Great Plains and the foothills of the Northern Rocky Mountains in southwest Montana. This range plant community was a more xeric form (gravelly soil) of the Palouse Prairie. Co-dominant range plant species were Idaho fescue and bluebunch wheatgrass. Green needlegrass and needle-and-thread were associate species.

These slides at three progressively (and distinctly) closer camera distances constituted a series of "nested photoplots". The first slide presented th physiogonomy (outward appearance) of this climax Pacific bunchgrass prairie while the second slide showed the physiogonomy at closer distance along with internal structure, including plant dispersion, of this range plant community. The third slide showed the internal structure including dispersion type or form, which was a random pattern, and the cespitose habit (morphological form) from a topdown view.

Students should note the complete absence of forbs from this steppe (bunchgrass prairie).

The stocking rate of cattle was on the light side of moderate so that degree of use was light with considerable herbaceous residue from last year's grass shoots.

The tufted or clumped habit of these bunchgrass or cespitose species was very pronounced; it was a textbook example to show Range Management students.

55. An enlarged cellulose cast- Sward of an Idaho fescue-bluebunch wheatgrass steppe on a livestock-big game range in part of the Palouse Prairie that developed in the foothills of the Northern Rocky Mountains. In the sample of range vegetation seen in these two "photoplots" green needlegrass and needle-and-thread were associate species on grassland that had almost no forb or shrub species. The larger green plants in the foreground of the second slide were green needlegrass and needle-and-thread.

Cellulose, which comprises much of the mature body of plants, is the most abundant organic compound on Earth. Much of the herbage (grass foliage) seen in the slides of this section was partly decomposed (rotted) organic matter (mostly cellulose) from last year's grass shoots. Such cellulosic material is low in forage digestibility so grazing animals instinctively avoid it and instead select actively growing plant material from this year's live grass shoots. (More high-cellulose herbage on this range was presented in the immeidately succeeding slides.)

56. Palouse Prairie pals- Idaho fescue and bluebunch wheatgrass, co-dominants of the Interior Northwest bunchgrass prairie or steppe that makes up most of the famed Palouse Prairie. These graminaceous folks were part of the climax range vegetation of a range grazed by beef cattle, mule deer, white-tailed deer, and elk. These plants had reached maximum development for this year's cool-growing season and had large panicles of ripening grain. This entire range was absolutely beautiful.

This cattle range (with mule and white-tailed deer and elk free-ranging on it) had been lightly stocked so that there weer still appreciable quantities of last year's grass shoots with the total herbage on offer. Grazing animals had largely avoided this high-cellulose, partly decomposed herbage.

Park County, Montana. Late June; early estival aspect with two co-dominant grasses at peak stading crop with ripening grain. FRES No. 36 (Mountain Grasslands Ecosystem. K-43 (Fescue-Wheatgrass). SRM 304 (Idaho Fescue-Bluebunch Wheatgrass). No biotic community designation in Brown et al. (1998). Range site: Foothills and Mountains- Silty, 15-19 inch precipitation zone (Ross and Hunter, 1976, p. 24). Northwestern Great Plains- Non-calcareous Foothill Grassland ecoregion 43s (Woods et al., 2002).

57. Slightly different neighborhood- In another local area of a climax Idaho fescue-bluebunch wheatgrass steppe in the Palouse Prairie plants of green needlegrass and needle-and-thread, the two associate species, made up a substantial portion of grass cover (both foliar and basal).

These three slides made up a set of "nested photoquadrants". The first slide presented physiogonomy (outward appearance) of this bunchgrass grassland while the second and third slide presented internal structure and composition of this steppe in which green needlegrass and needle-and-thread made a pronounced apperance in what was otherwise a "near half-and-half mixture" of bluebunch wheatgrass and Idaho fescue. The two needlegrass species were larger overall than the co-dominant grasses.

All four of these grass species are strictly cespitose (tufted or bunched) species. Idaho fescue, bluebunch wheatgrass, and needle-and-thread are on the Society for Range Management-sponsored International Intercollegiate Range Plant identifiction Contest (Stubbendieck et al., 1992, ps. 170-171, 188-189, 202-203, for Idaho fescue, needle-and-thread, bluebunch wheatgrass, respectively).

58. Don't forget us- Various species of lichen growing on rocks on an Idaho fescue-bluebunch wheatgrass range in the Palouse Prairie Region. This range was at the union of the Northern Great Plains and foothills of the Northern Rocky Mountains.

59. Hedged- A Rocky Mountain juniper (Juniperus scopulorum) hedged by deer (both mule deer [Odocoileus hemionus] and white-tailed deer [O. virginianus] are free-ranging in this area) on a foothill range grazed primarily by cattle in southwestern Montana on the extreme edge of the Northern Great Plains at the footslope of the Northern Rocky Mountains.

Park County, Montana. Late June.

Outstanding example of the bunchgrass prairie physogonomy of this range vegetation.

FRES No. 36 (Mountain Grassland Ecosystem. K-43 (Fescue-Wheatgrass). SRM 304. Northwestern Great Plains- Non-calcareous Foothills Grassland Ecoregion, 43s

(Woods et al., 2002). Park County, Montana. Late June (early summer); hard dough to mature phenological stage.

60. Some of the locals- Local population of Idaho fescue (with one or two "stray" plants of bluebunch wheatgrass) on the shallow, gravelly soil of the range featured above. A good example of the bunchgrass or cespitose habit of Idaho fescue.

FRES No. 36 (Mountain Grassland Ecosystem. K-43 (Fescue-Wheatgrass). SRM 304. Northwestern Great Plains- Non-calcareous Foothills Grassland Ecoregion, 43s (Woods et al., 2002). Park County, Montana. Late June (early summer); hard dough to mature phenological stage.

61. A few of the clumped folks- A closer-in view of some of the Idaho fescue plants on the foothill range featured here. The cespitose growth habit was shown. Also demostrated in this example were Idaho fescue plants that were less advanced in their maturity than most of the others. These plants had been grazed to a greater degree of use by beef cattle than most neighboring plants (though it was still light degree of utilization). This grazing defoliation appeared to have kept these plants in a less mature state of development (less advanced in the annual growth cycle). In fact, removal of the upper shoots had apparently kept--more-or-less--these individual in an asexual or vegetatively reproductive state.

Park County, Montana. Late June (early summer); pre-bloom stage of phenology.

62. A few that were a little farther along- Plants of Idaho fescue on the foothill range featured here that had advanced to almost the dormant state with spikelets of "hard seed". These plants were unusually large for Idaho fescue with several over two and a half feet in height.

Park County, Montana.

63. The slide was fine, but...- The Epson Perfection 600 scanner really "sparkled up" this very crisp image. Hopefully viewers can get some idea as to the size of some of the panicles of Idaho fescue from this example. This inflorescence, which was on one of the plants growing in the population presented in the last few slides, was almost two feet (two-thirds of a meter) in length. It was remarked above that Idaho fescue, a mid-grass species, has often been perceived as a comparatively small plant and yet this is frequently not the case at all, as in this instance. Total height in this example plant was almost a yard (meter) tall.

In this example, the actual panicle (not counting peduncle and lower sexual culm) was roughly two-thirds of total shoot height. For a perennial species Idaho fescue can produce an extraordinary yield of grain, tremendous allowation to sexual reproduction.

Park County, Montana. Late June (early summer); near grain-ripe phenological stage.

64. A particularly fine example- Robust plant of the cespitose bluebunch wheatgrass at peak anthesis. This larger-than-typical example of the wide-ranging bluebunch wheatgrass was growing on the sagebrush-bunchgrass savanna in the valley of the Little Big Horn River on the Northern Great Plains. This plant was accompanied--to its left and rear--by several plants of green needlegrass (Stipa viridula) which ranged from co-dominant to associate species to first-dominant, bluebunch wheatgrass.

Little Big Horn Battlefield National Monument, Big Horn County, Montana. Mid-June; peak standing crop at full anthesis.

65. Flowering on the fine example- Two spikes of the bluebunch wheatgrass specimen introduced immediately above at peak anthesis.

Little Big Horn Battlefield National Monument, Big Horn County, Montana. Mid-June; peak standing crop at full flower.

66. Bunched pollen on the Northern Great Plains- Spikelets of bluebunch wheatgrass at peak anthesis. This "photo-sample" was growing on the robust plant presented in the two immediately preceding images.

Little Big Horn Battlefield National Monument, Big Horn County, Montana. Mid-June; peak anthesis.43e Sagebrush Steppe

67. Bluebunch wheatgrass- The State Grass of Washington is the co-dominant (with Idaho fescue) of the Palouse Prairie and understories of ponderosa pine forests. Both species are locally abundant on some range sites of well-managed ranges in the Great Basin Desert. Bluebunch wheatgrass isone of the most valuable of western range plants but was largely displaced on many ranges by such exotic annual grasses as cheatgrass or downy brome and annual forbs like tansy mustard.

68. Inflorescences (spikes) of bluebunch wheatgrass- Rock Canyon, Utah County, Utah, June.

69. Population of Idaho fescue showing the cespitose habit of this bunchgrass. Idaho fescue is co-dominant with the less mesic bluebunch wheatgrass. Together these two valuable native forage species once dominated millions of acres of valuable range including grasslands like the Palouse Prairie to sagebrush-rabbitbrush shrub savanna steppe to the sparse herbaceous understory of the Great Basin desert. Early overgrazing on the unregulated open (ie. open to all comers; open range of the public domain) and establishment of alien annuals like cheatgrass caused loss of these two (and other native perennial grasses) from much of the Intermountain Region.

70. Idaho fescue (Festuca idahoensis)- This is the co-dominant of the Palouse Prairie or greater foothill prairie in the northern Rocky Mountain region. It is somewhat more mesic in habitat than it's co-dominant species, bluebunch wheatgrass.

71. Flower clusters of co-dominant bunchgrasses- Spike of bluebunch wheatgrass (left) and panicle of Idaho fescue (right) in the Middle Rocky Mountains. These two inflorescences were on plants that grew so close to each other that the photographer simply bent spike and panicle close enough together (and without damaging either) to snap these two shots.

Park County, Montana. Mid-June; caryopsis (grain)-ripening stage.

72. Another side-by-side slide of co-dominants- This was another growing and photographed side-by-side slide of bluebunch wheatgrass (left) and panicle of Idaho fescue (right). As was case with the preceding paired-comparison photographs, plants of the two species grew so close to each other that the photographer could "scoot" the two flower clusters this close together with harming either inflorescence.

This and the two preceding cheek-by-jowl, side-by-side slides illustrated the spatial closeness with which these two co-dominant species can grow over much of the Palouse Prairie. This seemed an appropriate juncture to remind students of the Competitive Exclusion Principle (no two species can occupy the same ecological niche or, said somewhat differently, one-species-one niche). To be consistent with this concept, one would have to argue that size and location of these two ecological niches were small and close together in the range vegetation of this "photo-sample" .

These two co-dominant species usually appear to get along quite well together on Palouse Prairie and ranges on adjoining areas. Perhaps faculty members and, even more so, faculty and administrators could take a lesson on thriving co-existence.

Sheridan County, Wyoming. Mid-June (late spring); soft-grain stage in bluebunch wheatgrass and anthesis in Idaho fescue. Next slide, please ...

73. Spreading joy and good cheer- Spikelets on part of a panicle of Idaho fescue at anthesis. These pollen-spreading organs were on part of the panicle shown in the immediately preceeding photograph.

Sheridan County, Wyoming. Mid-June (late spring).

74. Palouse Prairie pals in the foothills- Climax range vegetation of Interior Pacific bunchgrass prairie (an "island" of the Palouse Prairie) on a steep hillside in foothills of the Northern Rocky Mountains in southwestern Montana. Range plant species in the first "photoquadrant" (slide) included Idaho fescue, arrowleaf balsamroot (Bsalsamorhizas sagittata), one-flower false-sunflower or little sunflower (Helllanthella uniflora), sticky geranium (Geranium viscuissimus), and fringed sagebrush, prairie sagewort, arctic sage, and pasture sagewort (Artemisia frigida). The second "photoplot" (slide) featured Idaho fescue and fringed sagebrush or pasture sagewort and the sward they comprised.

The climax plant community on this hillside range was subject to grazing by white-tailed deer, mule deer, and elk. No livestock were on this range although it had undoubtedly grazed by domestic animals, especially sheep, in the past.

Park County, Montana. Mid-June; late vernal aspect. FRES No. 36 (Mountain Grasslands Ecosystem. K-43 (Fescue-Wheatgrass). SRM 304 (Idaho Fescue-Bluebunch Wheatgrass). No biotic community designation in Brown et al. (1998). Range site: Foothills and Mountains- Forest-Grassland Complex, Grassland Component thereof, 15-19 inch precipitation zone (Ross and Hunter,, 1976, p. 30). Middle Rockies- Mid-Elevation Sedimentary Mountains ecoregion 17g (Woods et al., 2002).

75. Two showy species- An "island" of Palouse Prairie In the foothills of the Northern Rocky Mountains furnished the habitat for two composites that are widespread and locally abundant in the grasslands and open forest of this region. One-flower false-sunflower or little sunflower (Helllanthella uniflora), two smaller plants at left background and left foreground, and arrowleaf balsamroot (Bsalsamorhizas sagittata), largest plant at center to right foreground, were in full-flower mode on this footslope grassland dominated by Idaho fescue.

Favorable soil moisture status made for nice "photogenic" specimens that were presented below.

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

76. Playing second fiddle to the big boy- One-flower little sunflower, one-flower false-sunflower, or little sunflower (Helllanthella uniflora) on a Northern Rocky Mountain foothill montane grassland dominated by Idaho fescue. This was a whole shoot view of the plant in left foreground in the immediately preceding slide. The reference to "little" in various common names for this composite are appropriate when most plants of this species are compared to such composites neighbors as arrowleaf balsamroot and mules-ears (Wyethia spp.). Otherwise it is a medium-sized forb.

One-flower false-sunflower is an important forb, at least in certain locales and local habitats, in the Palouse Prairie and parts of neighboring range plant communities, including open ponderosa pine forests. The Range Plant Handbook (Forest Service, 1940, W89) and Notes on Western Range Forbs (Dayton, 1966, p. 318) gave brief treatments of this member of the sunflower tribe (Heliantheae). One-flower false-sunflower has generally been rted as having Good forage value for sheep and deer and Fair forage value for cattle.

Park County, Montana. Mid-June.

77. An important forb of the Palouse Prairie- Arrowleaf balsam (Bsalsamorhizas sagittata) is perhaps the most characteristic, widespread, and showy composite, if not forb, on the Palouse Prairie and adjoining parts of neighboring range types such as ponderosa pine forests and savannahs, big sagebrush-bunchgrass shrub-steppe, and aspen parkland. The coverage in the standard reference, Range Plant Handbook (Forest Service, 1940, W43) and the brief treatment in the definitive Notes on Western Range Forbs (Dayton, 1966, p. 301) are good places for beginning Range Management students to introduce themselves to this important range forb. This valuable forb is on the list of 200 range plant species used in the International Intercollegiate Range Plant Identification Contest sponsored by the Society for Range Management (Stubbendieck et al., 1992, ps. 272-273). Another good reference for arrowleaf balsamroot was

There is general agreement among these sources as well asamong experienced rangemen in practice that arrowleaf balsamroot is good to excellent forage (depending on plant maturity) for small ruminants and fair for cattle and horses. This forb--and yet another member of the vast sunflower or daisy tribe (Heliantheae)--disappears from overgrazed ranges. It is a decreaser and telltale indicator species.

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

78. An important one- Perhaps no other forb so characterizes Palouse Prairie ranges in higher successional status tthan arrowleaf balsamroot. This forb species is also a member of the grassland communities of the more interior portions of coastal prairies of the Pacific Northwest. The species (biological) range of arrowleaf balsamroot extends into California and British Columbia.

Park County, Montana. Mid-June; full-bloom penological stage.

79. Montana had a lot choices, including this one- The Big Sky Country can rightly boast of having a number of "wildflowers" (native forbs) that would make fitting state flowers. Arrowleaf balsamroot even with the floral grandeur of the samplers seen here did not get the nod, but other states and provinces would be proud to have "wildflowers" just partly as eye-catching as this member of tribe Heliantheae. Examples seen here were growing on a steep, foothill slope dominated by Idaho fescue in the Northern Rocky Mountains. (It was a rough, slippery climb to bring back these trophies, but worth every step of it.)

Park County, Montana. Mid-June.

80. Breath-catching beauties- Capitula (heads) of arrowleaf balsamroot on a foothill montane grassland dominated by Idaho fescue in the Northern Rocky Mountains.

Park County, Montana. Mid-June.

81. Namesake of a major forb- Leaves of arrowleaf balsamroot. The specific epithet sagittata obviously refers to the very distinctive sagitate leaf shape. Well-named for an admirable range plant. This plant was growing in a grassland dominated by idaho fescue on a foothill slope in the Northern Rocky Mountains.

Park County, Montana. Mid-June.

82. One by itself- Capitulum of prairie or twin arnica (Arnica sororia) on a bluebunch wheatgrass-Idaho fescue grassland in the foothills of the Wyoming Basin.

Sheridan Countyk, Wyoming. Mid-June.

83. Phlox on the Palouse- Plains phlox (Phlox andicola) on a climax bluebunch wheatgrass-Idaho fescue bunchgrass steppe. This was the only forb (including the only plant of this species) that the author could find on this cow/calf range that was an island of Palouse Prairie in the Northern Rocky Mountains of southern Montana. This was as "pure" a grassland as this photographer ever saw: a "solid" bunchgrass prairie at climax stage, and this little forb was the only other notable herbaceous species that was not in the Gramineae.

Plains phlox is a rhizomatous, mat-forming species with short, often prostrate, shoots (Lesica, 2012, p. 395).

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

84. Fringed all over- Single plant of fringed sage or fringed sagebrush (Artemisia frigida) graowing on the same range beside the bluebunch wheatgrass and Idaho fescue on the outlier or island of Palouse Prairie in the Northern Rocky Mountains of southern Montana.

An excellent handy guide to the Artemisia species is the Montana Sagebrush Guide by Wambolt and Frisina (2002). Wambolt and Frisina (2002, p. 49) described fringed sagewort as a mat-forming species that occupies a variety of range sites and seral stages from pioneer to climax. The extremely soft material of this small shrub was used as bandages by Indians and frontiersmen. Dayton (1931, ps. 170-172) explained that browse value of this widely distributed species varied across its large species (biological) range, but that overall it was a locally important feed plant. In other areas it tended to be an invader, being common on overgrazed/overbrowsed ranges.

Park County, Montana. Mid-June; early pre-bloom stage.

85. Sparse and heavily cropped- A lone tree of Rocky Mountain juniper (Juniperus scopulorum) on an Idaho fescue-bluebunch wheatgrass foothill range being grazed by cow-calf pairs. Browsing was by mule deer (Odocoileus hemionus). Rocky Mountain juniper has the largest biological range of any Juniperus species in western North America (Burnsand Honkala, 1990). It is a shrub to small tree (Lesica, 2012) such that the specimen seen here was typical size and habit of Rocky Mountain juniper or Rocky Mountain cedar (the junipers are commonly referred to as "cedars" by stockmen, sportsmen, and the general populace.

The browse line on this plant was conspicuous.

Park County, Montana. Mid-June.

86. Rocky leader- Leader with fleshy seeds of Rocky Mountain juniper (Juniperus scopulorum) growing in the Northern Rocky Mountains. Leader is the term applied by foresters and rangemen to a woody branch. All Juniperus species are dioecious so this plant was obviously a female.

Gallatin County, Montana. Mid-June.

87. Rocky but fleshy- Fleshy seeds of Rocky Mountain juniper. Foliage and seeds shown here were growing in the Northern Rocky Mounatins. Two years are required for production and maturation of the seeds of Rocky Mountain juniper (Burns and Honkala, 1990). Fleshy seeds of Juniperus species are commonly called "berries", but of course conifers are not fruit-bearing plants having instead naked seeds usually referred to as cones. The "berries" of junipers are usually interpreted as fleshy cones. Jointed cone scales are visible on these fleshy seeds ((Lesica, 2012).

Gallatin County, Montana. Mid-June.

88. A woody one in an "island" of Palouse Prairie- Specimen of northern or fireberry hawthorn (Crataegus chrysocarpa= C. rotundifolia) growing on an island of bluebunch wheagrass-Idaho fescue in the moist dissected plains portion of the Northern Great Plains. This woody member of the rose family (Rosaceae), in the pear or apple subfamily (Pomoideae), has a species range from Newfoundland and Quebec westward to Idaho and Oregon southward to New Mexico and along the Atlantic Coast inward to Tennessee.

As a member of the pome subfamily of Rosaceae, the fruit of hawthorns is obviously of the pome type of fleshy fruits. (More details of this fruit was presented in the immediately succeeding slide-caption set where a cluster of hawthorn pomes was shown.)

Older, larger shoots shown in the dreafullly messed up (by an Epson Perfection 700 scanner) first slide appeared to have been topkilled by a wildfire two or three growing seasons previous. Northern hawthorn is apparently adapted to such fires . It probably increased number of shoots and total plant cover in response to this fire and loss of apical dominance by the large (now dead) shoot.

Sheridan County, Wyoming, Mid-June; immature-fruit phenological stage.

89. Cock spurs, rough edges, and drupes- Distal end of lateral leader (first or upper slide) and leaves and fruit cluster (second or lower slide) of northern or fireball ahwthorn growing on a bluebunch wheatgrass-Idaho fescue island of Palouse Prairie in the moist dissected plains portion of the Northern Great Plains. The hawthorns--as evidenced by their common name--are well known for their long thorns (often called "cock spurs" by cock-fighting hillbillies) and for their oval- to arsuate- to spatulate- shaped, serrate (sometimes lobe-like) leaves.

With possible exception of Carex among the monocots, Crataegus is the most difficult and confusing vascular plant genera in North America. Expert botanists in genera like these are regarded as the "brain surgeon-scientists" or "rocket- scientist" among taxonomists. This is due to the numerous recognized--and very similar--species of Crataegus plus hybridization among these species. For example, Steyermark (1963, ps.) gave 37 species of Crateagus for Missouri while the Great Plains Flora Association (1986, ps, 370-375) described 11 Crataegus species for much of the Great Plains and adjoining portions of the Central Lowlands provinces. Good soruces for C. chrysocarpa included Stephens (1973, ps. 234-235) and Turner and Kuhlmann (2014, p. 239).

Sheridan County, Wyoming, Mid-June; immature-fruit phenological stage.

90. Rough edges and shaggy base-Two boles (trunks, central shoots) of fireball hawthorn on an island of Palouse Prairie dominated by bluebunch wheatgrass and Idaho fescue in the mesic dissected portion of the Northern Great Plains. This Crataegus species often forms thickets as well as being found as a single shrub or small tree (Stephens, 1973, ps. 234-235).

Sheridan County, Wyoming, Mid-June.

There are many--too damn many--alien or exotic plant species that were introduced (most inadverdently) into North America by the white man. Many of these species naturalized to some extent or the other and became (now) nautrally occurring weeds, some more serious pests than others of course. Some of these weeds (weeds by any and all definitions, both economic and ecological meanings) are more or less strictly agronomic or horticultural pests that require farming-type disturbances (eg. tillage, planting, intense harvest) to perpetuate their co-evolved-with-humans existence. These agronomic-horticultural weeds are of little or no moment in context of Range Management. Other alien plants adapted so completely to their new home that they appear to be "thoroughly" naturalized and are now range plants the same as if they were natives ("original" North American range plants).

Such noxious vascular range plants include both woody species (that are properly labeled as brush) and herbaceous species that are by convention properly called weeds in the more restrictive definition of this term. Some of the latter (= herbaceous noxious plants) have minor influences in range plant communities whereas others have major--often all-determining-- ecological impacts on range vegetation (frequently such disruptive affects as to threaten or even collapse range ecosystems). This results in consequent adverse socioeconomic consequences on human society. Other weeds (herbaceous pests) have a minor impact on man and Nature resulting in some positive as well as negative effects. The status of weed or weediness for these exotic plants varies depending on prevailing conditions and, consequently, from one livestock operation, wildlife refuge, commercial forest, and even one range site to the next. Degree or extent of noxiousness can vary year-to-year depending not only on density and cover of weeds, but also on benefits or costs to the individual management unit. For instance, weeds have saved many animals from starvation, ranchers from bankruptcy (at least from a few light meals), and acres of otherwise bare land in drought, recession, and other "storms" of one kind or the other.

Some of these "good news-bad news" naturalized weeds of the Palouse Prairie (and adjoining areas) were included in the immediately following portion. Others were treated above and below this segment so as to be covered with the range plant communities in which they were photographed.

91. Bromus tectorum- This naturalized Eurasian annual known variously as cheatgrass, downy brome, and downy chess is probably the Bromus species most widely distributed and dominant over the most area in North America. Pavlick (1995, p. 7) recognized 20 or 21 annual non-native Bromus species in North America, many of which "are weedy species that occupy disturbed sites such as fields, waste places, road verges and overgrazed rangeland" ((Pavlick, 1995, p. 8). B. tectorum falls in that category. It is in the section Genea which includes such other introduced and naturalized annual bromegrasses as ripgut brome (B. rigidus), great brome (B. diandrus), and red brome (B. rubens). Cheatgrass-dominated ranges usually have not been regarded as range cover types, or at least not desirable range types. The Society for Range Management (Shiflet, 1994) did not recognize the disclimax of cheatgrass range as a cover type. This differs from the case of the California annual grassland in which several of the Genea section bromes are dominant and associate species. While the California annual type is a disturbance climax, it is a species-rich grassland or oak- annual grass savanna and it is accepted as a rangeland cover type which is now the potential plant community and the range plant community for which to manage. By comparison, cheatgrass is not only a disclimax but in effect an accidental man-made "monoculture" which is usually viewed as a more or less undesirable state of range retrogression. Unlike some of the naturalized Eurasian annual bromes in the California annual type, which respond as and are interpreted as decreasers or increasers (eg. ripgut and soft brome), cheatgrass is a classic invader. Furthermore, Bromus tectorum does have the forage value (palatabability and nutritive content) of Eurasian annuals like B. rigidus or B. mollis.

Most importantly, downy brome is a dreadful weed because it dominates tens to hundreds of thousands of acres of Intermountain rangeland that have the potential to return through secondary succession to something resembling the more productive and nutritionally superior pre-Columbian climax grassland. At least, native perennial bunchgrasses like bluebunch wheatgrass, Idaho fescue, and Sandberg bluegrass could increase and become more abundant than they are where it not for fierce competition with cheatgrass. Cheatgrass initiates growth earlier in the spring than the native perennials and depletes soil moisture thereby depriving the native bunchgrasses of the most limiting abiotic factor in the range ecosystem. This is most pronounced in seedling development because cheatgrass seedlings grow faster and extend their roots deeper earlier in the winter than do seedlings of the native perennial grasses (Harris, 1967).

Another major factor responsible for the persistence of cheatgrass (and deteriorated range) is the role of dead cheatgrass residue as a source of fuel for lightening-ignited fires later in the summer fire season. Cheatgrass completes its life cycle earlier in the growing season than the native perennial grasses and when the straw of the prolific seed-producing annual grass burns this damages the natives which are still growing and which rely to a larger extent on asexual reproduction by tiller production. With this combination of factors and conditions (and perhaps others like competition for light or selective grazing of the more palatable but less grazing-tolerant bunchgrasses) cheatgrass perpetuates it's ecologically and nutritionally inferior "monoculture". The reason why the perennials are more nutritious over a longer period is that they take longer to complete their annual growth cycle. This is what is meant by a longer green feed season. The shorter or faster growth cycle of annual grasses allows the continued displacement of the native perennial grasses by annual alien weeds. This results in reduced forage value and lowered quality of range livestock and wildlife diets.

As is the case with any higher plant cheatgrass, including its residue of dead straw or mulch, provides certain benefits to range and meets some requirements for range ecosystem function including a degree of protection against soil erosion and watershed degradation, addition of organic matter to soil, source of forage and other habitat factors for wildlife (eg. nesting material for birds), sequestering of carbon, etc. Cheatgrass even has a subjective but distinctive beauty as its fuzzy shoots and lacy panicles grow and change colors with progression of season. In these ways, downy brome does fulfill certain required features or properties necessary for "rangeland health" (Committee on Rangeland Classification, 1994; Natural Resources Conservation Service, 1997, ps. 4-23-4-26; Mitchell, 2000, ps. 27-55), whatever the heck "rangeland health" is.

Among other things, these benefits of cheatgrass (again, any vascular plant species furnishes such benefits) allows managers of public land to assert that depleted cheatgrass range is not as bad as might appear strictly on basis of traditional range condition/trend analysis because cheatgrass cover does offer enough of the things just listed to prevent further deterioration of natural resources. This means that replacement of native perennials by cheatgrass permits degradation of range ecosystems to progress only to the point that basic resources like soil and water are not extremely, at least not totally, exhausted). In other words, cheatgrass populations can, on some range sites and under certain situations, sustain "the integrity of the soil and the ecological processes of rangeland ecosystems" enough to meet "the minimum standard"…"to prevent human-induced loss" or "rangeland health" (Committee on Rangeland Health, 1994, ps. 4-5).

A specific, if hypothetical, example might be a climax shrub steppe community of bluebunch wheatgrass, Idaho fescue, bitterbrush, and big sagebrush that through overgrazing a century ago and uncontrolled summer wild fires in recent times deteriorated to a "solid stand"of cheatgrass. By previous standards of range condition and trend based on successional state (seral stage) this range was (or would have been) rated as Poor or Very Poor, it being at one of the lowest seral stages due to retrogression. Now not to worry: based on the revised standards of "rangeland health" the range ecosystem could be in or at an acceptable ecological status because the soil is not washing away and the stream draining the watershed is not carrying an unacceptable sediment load. Cheatgrass ("any plant") recycles nutrients and acts as a factor in soil formation. In short, any plant (even an alien weed like cheatgrass) contributes to ecosystem structure and function and, thus, "rangeland health".

Furthermore, it might be plausible that a cheatgrass-dominanted range community could be the "desired plant composition" for certain uses and values (Committee on Rangeland Health, 1994, p. 94) "at least for a period of time" based on standards of "soil stability and watershed function, integrity of nutririent cycles and energy flow, and presence of functioning recovery mechanisms" (Committee on Rangeland Health, 1994, ps. 98-99). This does not mean that the land or plant community has to have recovered, but only that recovery mechanisms "must be in place and they must be working" as determined by the criterion of "changes in plant demographics" (Committee on Rangeland Health, 1994, p. 120). It was not clear in examples given by the Natural Resources Conservation Service (1997, ps. 4-23-4-26) if and for how long some cheatgrass ranges would be in an acceptable state of "rangeland health" if there were not "changes in plant demographics". Nor was it clear the spceific meaning of these "demographics". For instance, does this mean the traditional criteria of plant succession such that over time the same parcel of land comes to be occupied by plants of higher ecological (= successional) status (eg. gradual replacement of cheatgrass by decreasers like bluebunch wheatgrass)? Or, alternatively, can "demographics" apply at the population level such that an increasing number or cover of cheatgrass over time is a change in "demographics"?

It would appear that Bromus tectorum is fodder for thought as well as for range animals and fires. By the way, when cheatgrass serves as fuel to increase frequency of wild fires "hereby leaving soils exposed to erosion and watersheds more apt to degrade" how is this factored into the calculation of "rangeland health"? Erath County, Texas. April.

92. Cheatgrass panicles- This mass of spikelets shows the nature and extent of sexual reproduction in annual grasses. This is an example of the therophyte life- or growth-form category of Raunkiaer. Therophytes are annual or ephemeral plants which are generally the life-form most tolerant of conditions marginal for life because they complete their life cycle rapidly and spend most of their life cycle as the gametophyte generation (ie. seeds). In the Raunkiaer life-form concept the therophyte category is the general plant group best protected from such conditions as drought, extreme temperatures, and such forms of defoliation as grazing, fire, wind, frost, hail, etc. Perennial grasses generally are in the cryptophyte or hemicryptophyte life-form whose perennating buds and shoot apices are fairly well protecting by being at or below the soil surface. This is, of course, why defoliation by grazing, fire, desiccating winds, etc. tend to damage these life-forms less than the phanerophyte life-form. The net result being that such stresses favor directly (and indirectly through competitive advantage) those plants with lower meristematic tissues, other things being approximately equal (eg. palatability and flammability of plant parts).

Under the most severe extremes of grazing/browsing degrees of use, harsh fire regimes, drought, and desert climates therophytes become progressively more common or the dominant life-form. Under (and often for decades following) prolonged and severe overgrazing, protracted drought, and repeated fire at critical stages of plant phenology therophytes come to dominate the other life-forms, including even the cryptophytes and hemicryptophytes, all the more. Thus throughout the Intermountain Region between the Rockys and the Sierra Nevada-Cascade ranges cheatgrass came to dominate and displace, often to a monoculture-like exclusion, the native perennial bunchgrasses, valuable browse plants (eg. antelope bitterbrush), and even the relatively unpalatable shrubs like sagebrush, rabbitbrush, and greasewood.

Erath County, Texas. April.

The Society for Range Management (Shiflet, 1994) did not recognize cheatgrass range as a rangeland cover type and this author agreed with that omission. Palouse praire and big sagebrush shrub steppe that has deteriorated is not in this author's judgment a "neoclimax" or disclimax (a anthropogenic climax) that is to be managed as if it was native. That is the situation for California annual grassland, but not for cheatgrass-mustard range. The California annual type is a botanically diverse community in which the naturalized Mediterranean annual grasses and forbs even respond in the classic decreaser-increaser-invader model of range retrogression. The forage value of many of these Eurasian species is often greater while they are growing than for the native bunchgrasses. These situations do not obtain for weedy annuals in the Intermountain Region. Cheatgrass forms a "monoculture" not a diverse community of numerous species and niches. Weeds of the Great Basin and Palouse Prairie were included here for the purpose of showing students textbook examples of invader species.

For currently comprehensive coverage of the problem of cheatgrass and other associated weeds on Intermountain ranges see McArthur et al. (1990) and Monsen and Kitchen (1994).

Some of the most important naturalized forbs that are weeds in the Palouse Prairie (and Intermountain Region) are crucifers, members of Cruciferae (the mustard family). These include annual Eurasian species.

93. Pinnate tansy mustard or tansymu;stard (Descurainia pinnata= Sisymbrium pinnatum)- This is a characteristic population of one of the most thoroughly naturalized and invasive weeds on the western range. This Eurasian annual crucifer is a dreadful weedy forb that in association with cheatgrass or downy brome has displaced the native bunchgrasses and browse plants throughout the Intermountain Region. Tansymustard has naturalized across most of the North American continent, but it is across the Western Range where it grows in dense populations such as seen here so as to constitute a disutrbance climax that is probably a permanent range cover type, at least in human time scale. In immature stages tansy mustard can be fair to good forage for small ruminants, but it is often toxic to cattle in which it causes tongue paralysis and inability to swallow even water.

The Society for Range Management designated this naturalized forb as one of its "200 list" for the International Intercollegiate Range Plant Contest (Stubbendieck et al., 1992, ps. 292-293).

Wasatch County, Utah. June

94. Flowers of tansy mustard- Members of Descurainia are generically known as tansy mustards. Taxonomic treatment of several of the cruciferae genera has presented workers with problems over the years and some of these species have placed in various genera over the years. Wasatch County, Utah. June.

95. Flixweed (Descuriana sophia)- Upper shoot of another Eurasian winter annual crucifer that naturalized to become a weed on Palouse Prairie range. Siques were nearing maturity on this specimen and fairly conspicuous. Flixweed is much less widespread and only a minor weed on Palouse Prairie range. It often grows in asociation with pinnate tansymustard and other weeds. The one shown here was growing alongside tall tumble mustard and other weeds.

Latah County, Idaho. J:une (early summer).

96. Tall tumble mustard or tumblemustard (Sisymbrium altissimum)- Along with the tansymustards, especially pinnate tansy mustard, the tumblemustards are the some of the most important, Eurasian, weedy forbs in the Palouse Prairie and generally over the Intermountain Region. A distinguishing feature of tall tansymustard is the distinctive two forms of lower and upper leaves. Lower leaves in this plant that was growing in extremely dry soil had already been shed negating that feature in this specimen, but characteristic finely divided upper leaves (that contrast with wide or broad lower leaves) were obvious.

Latah County, Idaho. June (early summer); floweing in upper portions of inflorescence while approaching mature fruit stage in older siliques.

97. Shoot of tall tumblemustard- Two views of upper shoot of Sisymbrium altissimum showing details of upper leaves, fruit, and infloresecence. In tall tumblemustard pedicels of siliques are considerably thicker or larger in diameter than siliques themselves.

Latah County, Idaho. June (early summer); still floweing but also approaching mature fruit stage in lower (older) silques.

98. Flowers of tall tumble mustard- Details of the inflorescence of this widespread weedy forb. Latah County, Idaho. June (early summer); peak bloom in upper parts of inflorescence and approaching fruit maturity of older silques.

99. Small tumblemustard or little tumbling mustard (Sisymbrium loeselii)- This species closely resembles tall tumblemustard at first glance, but small tumble mustard has the same basic leaf shape throughout the plant. Also the pedicels of siliques are noticeably smaller in diameter than diameter of mature siliques. Furthermore, as suggested by their common names, small tumblemustard is smaller than tall tumblemustard (maximum heights of two vs. five feet). S. loeselii is much less important (other than at local scale) on range than is tall tumblemustard. Given ease of confusion between these two similar species and the importance of S. altissimum as a common range weed it was deemed appropriate to include S. loeselii at this juncture.

Okanogan County, Washington. June (early summer); full-bloom stage with lower silques ripening

100. Basal leaves of small tumblemustard- Close-up photograph of the above specimen of S. loeselii showing characteristic lower leaves of this species. Okanogan County, Washington. June (early summer).

101. Sexual shoot of small tumblemustard- General view of inflorescence with lower units having formed fruit (siliques) while upper units in this indeterminate-flowering species are still in peak bloom. Silique is a fruit type that is elongate and dehisces lengthwise (along sides of elongated "pod-like" fruit) leaving the internal partition called the replu;m (Smith 1977,p. 307).

Okanogan County, Washington. June (early summer).

Channeled Scablands Range

"Aside from affording a scanty pasturage, scabland is almost without value".-- J. Harlen Bretz

It was explained in the introduction to the bunchgrass prairie (a steppe dominated by cespitose grasses) that this formation or subformation (depending on interpretation of formation) is--at least was in pre-Columbian North America--in contact with the mixed prairie-shortgrass plains grasslands to the east. Clements (1920, p. 150) used as incontrovertable ecological evidence of this the presence of needle-and-threae (Stipa comata), Junegrass (Koleria cristata), and western wheatgrass (Agropyron glaucum= A. smithii) in both grasslands (grassland formations or subformations). Likewise the bunchgrass prairie of the Interior Pacific Northwest is in contact with the bunchgrass prairie of California or the Pacific Slope (Coastal Pacific Northwest) as indicated by presence of Idaho fescue (Festuca idahoensis) in both these grassland units.

Palouse Prairie has traditionally been the proper noun used as a synonym for the interior bunchgrass prairie, a regional grassland whose natural boundaries extended far eastward of the domain of the Palouse (Palus) Indian tribe, the namesake of this distinctive steppe. This usage was applied in particular by early workers like F.E. Clements and J.E. Weaver and later students such as H.F.Heady so that the traditional name persisted to remain in wide and common usage-- poetic, geologic, and ecological as well as commercial.

As interpreted by the current author the Palouse Prairie Region includes grasslands (steppes) that developed on land within the distinctive land form known as channeled scablands. This land is closer in proximity and geologic history to the Palouse Hills of the Rocky Mountains than this latter land form is to the eastern margins of the Palouse Prairie bunchgrass ranges on the Great Plains. Often in contradistinction to the deep, fertile soils in those parts of the Palouse Prairie within the Northern Rocky Mountains and along the eastern Columbia Plateau, the scablands (as indicated by that descriptive title) are frequently shallow, rocky, and dust-dry in some "spots" yet deep and wet in adjacent depressions. Rock outcrops and cistern-like piles of igneous rock often exist in between these extremes. Such edaphic-topographic variation at relatively small spatial scale renders variation in range plant communities over a given area of scablands among the greatest on the continent. In fact, differences in range sites and/or habitat types (hence, range vegetation) on channeled scablands are perhaps exceeded only by those on mountain sides and, especially, mountain tops (ie. alpine range ecosystems, plant communities, range sites, etc.).

Channeled scablands featured in this publication were in the Walla Walla section (20a) of the Columbia Plateau physiographic province (Fenneman, 1931, ps. 251-252). Formation of channeled scablands (ie. their geologic history) and the discovery of the geologic forces responsible for their formation (ie. history of their discovery and explanation) remains one of the most intriguing in American geologic circles. It was beyond bounds of the present work to give geologically detailed accounts of scablands. Readers were referred to standard texts and references for such. This author's favorite was the account devoted exclusively to this geologic wonder (especially its eventual explanation), Cataclysms on the Columbia (Allen et al., 1986). The channeled scablands of the Columbia Plateau not only resulted in some of the most interesting and locally variable range vegetation on the Western Range, but their final explanation by geologist J.H. Bretz was as cataclysmic to the discipline of Geology as the floods he described were to the land and water resources of the region.

Simply put, about 20,000 years BP a big glacier reached that part of the Interior Pacific Northwest that is today parts of Montana, Idaho, and Washington. Glacial ice formed a dam and created Lake Missoula. When the ice dam melted, shifted, or whatever it broke therby releasing, it has been estimated, more than a hundred times as much water as in the Mississippi River. This resulting cataclysmic flood, the Spokane Flood which was perhaps the greatest river flood in Earth history, scoured out the channeled scablands. Aerial views of the flood's course and maps of the channeled scablands it formed resemble the pattern of a great braided stream. Rushing flood waters carved some parts of the land deep into the basaltic parent material while leaving other close-by parts of the land much less scoured. More precisely, there was not just one flood but perhaps as many as 40 such cataclysmic floods (often called Bretz floods) over parts of the Columbia River drainage basin during a span of roughly two millennia (2000 years) somewhere in time between 12,000 to 15,000 years ago (Allen et al., 1986, p. 103).

This explanation by J.H. Bretz was for many years rejected by the geologic "establishment" for, among other things, it revived the debate between the two general--and opposing--doctrines used to explain certain geologic pheomena: Uniformitarianism and the older or original Catastrophism. Bretz' explanation of the Spokane Flood and channeled scablands was eventually accepted (later near the end of his long life) and proved that some geologic events are indeed cataclysmic.

From the point of view of Range Management the "heap big" flood carved out some of the most scenic, diverse, and interesting range to be found. From a rangeman's view of the world one of the best parts about the channeled scablands is that ain't nobody going to stick a bloody plow in their rangeland.

A recommended summary of soil formation and soils in general of the western Palouse Prairie and adjoining channeled scablands was the discussion of H.R. Gentry in the Soil Survey of Whitman county, Washington (Soil Conservation Service, 1980, ps. 149-173 passim).

The one rangeland cover type described by the Society for Range Management (Shiflet, 1994) that is found in the channeled scablands was Bluegrass Scabland (106). That title and its description certainly covered the Sandberg bluegrass-dominated range in the scablands and which are a major part of the channeled scablands complex. Likewise SRM Types 101 (Bluebunch Wheatgrass), 102 (Idaho Fescue), and 302 (Bluebunch Wheatgrass-Sandberg Bluegrass), along with some others perhaps, cover much of the remaining kinds of range (range types, range sites, habitat types depending on spatial scale, sucessional interpretation, etc.). However, these descriptions (including that of Bluegrass Scabland) did not make it clear to the reader who had yet to visit the channeled scablands that these were part of that specific land form nor that the scablands were much more encompassing than Bluegrass Scabland (SRM 106). Neither did the Society for Range Management (Shiflet, 1994) include the sporadic, but widely distributed shrub garlands on rock outcrops, igneous talus, etc. (Franklin and Dryness, 1973, ps, 229-230) on channeled scablands. Most confusing and deficient of all was absence of any rangeland cover type for basin wildrye-dominated rangeland or for marshes that are within and part of the channeled scabland complex. Given absence of designations and descriptions of these range types, the coverage of channeled scablands offered below of necessity was lacking appropriate names or titles of range types within Columbia Plateau channeled scablands. With students duly advised of this deficiency coverage began.

The first example of channeled scablands range was of various climax range plant communities in the Channeled Scablands Ecoregion (10a) of the Columbia Plateau that was adjacent to and just west of the Spokane Valley Outwash Plains Ecoregion (15s) of the Northern Rocky Mountains and north to northwest of the Palouse Hills Ecoregion (10h) of the Columbia Plateau.The following photographs of channeled scablands range vegetation includes those range sites or range subtypes of the Palouse Prairie steppe of the Agropyron spicatum-Poa sandbergii (= P. secunda) zonal association of Franklin and Dryness (1973, p. 211). This unit of range vegetation regarded is part of Society for Range Management (Shiflet, 1994) rangeland cover type 106 (Bluegrass Scabland) or, alternatively, SRM cover type 106 can be interpreted as one of several range sites, habitat types, or range subtypes found within the land form of channeled scablands. Other major, native, perennial grass species besides bluebunch wheatgrass and Sandberg's bluegrass included basin wildrye, squirreltail bottlebrush (Sitanion hystrix), Thurber's needlagrass (Stipa thurberiana) and, less commonly, needle-and-thread (S. comata). Less common still were the naturalized, Eurasian, annual grasses: cheatgrass or downy brome and bulbous bluegrass (Poa bulbosa).

Most important forbs (in approximate order of commonness or relative abundance which was a general combination of cover and density based on occular estimate) were: narrowleaf skeleton weed or bush wire-lettuce (Stephanomeria tenuifolia var. tenuifolia), western yarrow (Achillea millefolium ssp. lanulosa), Gorman's lomatium (Lomatium gormanii), silky lupine (Lupina sericeus), and blanket flower (Gaillardia aristata), and pale agoseris (Agoseris glauca var. glauca). Major shrubs included basin big sagebrush and stiff or scabland sagebrush (Artemisia rigida), alternatively the dominant woody species, rubber rabbitbrush (Chrysothamnus nauseosus), a sometimes local dominant or co-dominant woody species, snow wild-buckwheat (Eriogonum niveum) and creamy, parsnip-flowered, or Wythe's wild-buckwheat (E. heracleoides).

102. Selected views of channeled scablands range of bluebunch wheatgrass-Sandberg bluegrass- Four photographs of a drier form of bunchgrass steppe on Columbia Basin scablands with various local habitats dominated variously by Sandberg bluegrass, bluebunch wheatgrass, stiff sagebrush, basin big sagebrush, and basin wildrye. This range had been grazed by cattle and mule deer. Cattle had been moved to other range. The first of these four slides was typical Sandberg bluegrass dominated range with bluebunch wheatgrass either weaker dominant or associate species. Cheatgrass and, at much lower densities, bulbous bluegrass were present.

The second of these photographs was atop an igneous shallow scab topographic feature (sometimes called "biscuit scab") with basin big sagebrush dominant or co-dominant with bluebunch wheatgrass and Sandberg's bluegrass in some microhaibtats and with . The third slide was of range vegetation just below or off of the shallow scab. Foreground vegetation was dominated by scabland or stiff sagebrush but, which like that in the second slide, was of smaller size than would be the case on deeper soil with sparse cover of Sandberg bluegrass, cheatgrass, and bulbous bluegrass. Range vegetation in midground was combination of these same three grass species plus bluebunch wheatgrass along with heavily grazed narrowleaf skeleton weed or bush wire-lettuce. Immediately behind there was a microsite comprised of a small pothole with deeper soil in which basin wildrye made up most of the plant community. Western yarrow, silky lupine, and blanket flower were also present along with some Sandberg bluegrass and the two Eurasian annual grasses. The fourth photograph in this series of four was a closer-in, detailed view at edge of the basin wildrye-dominated microsite. Silky lupine, western yarrow, blanket flower and basin wildrye were readily visible as was the dull rust-colored dormant herbage of cheatgrass, Sandberg bluegrass, and bubous bluegrass.

While this range vegetation may have looked "purty sparse" to newcomers to these parts this range was almost assuredly in Good range condition class based on species composition of the existing vegetation compared to climax range vegetation. This range had been grazed by cattle and overall it was a relatively dry habitat even by scablands standards so it looked "rough", but degree of use was not excessive (it probably fell within the bounds of proper use) and the range plant community was dominated by the climax dominants, Sandberg bluegrass and bluebunch wheatgrass, and not by Eurasian annual grasses (invaders).

Lincoln County, Washington. June (estival aspect) Dormancy and grain-shatter stage in Sandberg's bluegrass, bulbous bluegrass, and cheatgrasss; peak standing crop and hard-dough stage in bluebunch wheatgrass; full-bloom stage in the three forbs. FRES No. 36 (Mountain Grasslands Ecosystem). K-44 (Wheatgrass-Bluegrass). SRM rangeland cover type designation would include primarily SRM 106 (Bluegrass Scabland), but also some areas would be better described as SRM 302 (Bluebunch Wheatgrass-Sandberg Bluegrass). This latter was the Agropyron spicatum-Poa secunda habitat type, most of which was the lithosolic phase (Daubenmire, 1968, ps. iii, 47-49). Wheatgrass Series of or within Great Basin Shrub-Grassland biotic community or regional formation of Brown et al. (1998), but there should be a Bluegrass Series under that and some of this range vegetation would be of that designation. Soils were primarily of the Anders-Bakeoven-Rock Outcrop Complex mapping unit; range sites for these two series were Loamy (L-2) and Very Shallow, respectively (Soil Conservation Service, 1981, ps. 11, 108, 110). The Very Shallow range site (Bakeover series) clearly predominated. Columbia Plateau- Channeled Scablands Ecosystem, 10a (Environmental Protection Agency, undated).

103. Inside the partially healed wounds- J. Harlen Bretz described the scablands as "wounds only partially healed" (Allen et al., 1986, p. 31). The first of these two slides presented the general topographic features along ridge tops or or crowns of the highland parts of channeled scablands showing numerous outcroppings of igneous rock as well as "scabs" or "scalds" existing where soils are very shallow to non-existant. One of these shallow-soil microsites was present in midground at left margin. The second of these slides was of the "scald" (thin-soil microsite). Sandberg's bluegrass was the ominant range plant in vegetation presented in both slides. Scrubby individuals of stiff or scabland sagebrush dominated much of the otherwise bare land of the "scab". Much of the darker, reddish straw (as in background of both photographs) was cheatgrass. Bluebunch wheatgrass was conspicuous as the only green plant present aside from sagebrush and an occasional western yarrow.

Lincoln County, Washington. June (estival aspect) Dormancy and grain-shatter stage in Sandberg's bluegrass, bulbous bluegrass, and cheatgrasss; peak standing crop and hard-dough stage in bluebunch wheatgrass; full-bloom stage in the three forbs. FRES No. 36 (Mountain Grasslands Ecosystem). K-44 (Wheatgrass-Bluegrass). SRM rangeland cover type designation was almost exclusively SRM 106 (Bluegrass Scabland), but also some areas would be more completely described as the Artemisia rigida-Poa secunda habitat type (Daubenmire, 1968, ps. iv, 40-42). Wheatgrass Series of or within Great Basin Shrub-Grassland biotic community or regional formation of Brown et al. (1998), but there should be a Bluegrass Series under that and some of this range vegetation would be of that designation. Soil was mostly of the Anders-Bakeoven-Rock Outcrop Complex mapping unit, but mostly Bakeoven series the range site of which was Very Shallow (Soil Conservation Service, 1981, ps.11, 108, 110). Columbia Plateau- Channeled Scablands Ecosystem, 10a (Environmental Protection Agency, undated).

104. View of and on a scab- On another range, adjacent to the range presented immediately above, the channeled scablands had different range sites--hence, different range plant communities--and a different range condition class. The range vegetation shown in these two slides featured a so-called biscuit scab: a side-view of it in the first slide and a top-view of it in the second. A range examiner's view of this rangeland vegetation revealed a range in Fair range condition class in contrast to Good condition class of the previously "examined" (immediately above) channeled scablands range. While the range being examined in these photographs had more grass cover this grass was mostly cheatgrass rather than the climax dominants of Sandberg bluegrass and bluebunch wheatgrass (ie. this range was generally dominated by an Eurasian invader). Soil was deeper on this range than on the range viewed in the preceding set of photographs such that this range (this combination of range sites) had higher production potential for grass biomass, and for that standing crop to come from the climatic climax Agropyron spicatum-Poa sandbergii (= P. secunda) zonal association (Franklin and Dryness, 1973, p. 211). Instead, this range was in a state of retrogression, essentially a disclimax of Bromus tectorum. While this channeled scablands range had "more grass" it was the wrong species of grass based on ecological or potential natural vegetation.

In the first photograph there were some plants of bluebunch wheatgrass (visible as green clumps) and even more of Sandberg's bluegrass, but these were overwhelmed by cheatgrass. In this slide the short shrub in right foreground (also represented by some plants barely visible in background) was creamy, parnship-flowered, or Wythe's wild-buckwheat. A biscuit scab was visible from the side-veiw in left to center background. The second slide was taken atop the biscuit scab showing another example of the Artemisia rigida-Poa secunda habitat type (Daubenmire, 1968, ps. iv, 40-42) introduced above in foreground. (Sandberg's bluegrass was better represented than scabland sagebrush in this sample of that habitat type.) Larger patches of green foliage in midground and distant background were stands of basin wildrye.

Lincoln County, Washington. June (estival aspect) Dormancy and grain-shatter stage in Sandberg's bluegrass and cheatgrasss; peak standing crop and hard-dough stage in bluebunch wheatgrass; full-bloom stage in forbs. FRES No. 36 (Mountain Grasslands Ecosystem). K-44 (Wheatgrass-Bluegrass). SRM rangeland cover type designation was almost exclusively SRM 302 (Bluebunch Wheatgrass-Sandberg Bluegrass) which is the climatic climax Agropyron spicatum-Poa sandbergii (= P. secunda) zonal association of Franklin and Dryness (1973, p. 211) and also the the Agropyron spicatum-Poa secunda habitat type (Daubenmire, 1968, ps. iii, 18-20), which was in contrast to the lithosolic phase of this habitat type featured in the scablands range presented previously. This range was in some degree of degradation with much of the vegetation being a cheatgrass disclimax. Local areas of the range shown here (eg. top of biscuit scab) were the edaphic climax Artemisia rigida-Poa secunda habitat type (Daubenmire, 1968, ps. iv, 40-42). Wheatgrass Series of or within Great Basin Shrub-Grassland biotic community or regional formation of Brown et al. (1998). Soils were primarily of the Anders-Bakeoven-Rock Outcrop Complex mapping unit; range sites for these two series were Loamy (L-2) and Very Shallow, respectively (Soil Conservation Service, 1981, ps. 11, 108, 110). The Very Shallow range site (Bakeover series) clearly predominated on the biscuit scab whereas the Loamy (L-2) range site was the more widespread range site across the surrounding rangeland. Columbia Plateau- Channeled Scablands Ecosystem, 10a (Environmental Protection Agency, undated).

105. Pretty is as pretty does- Channeled scablands rangeland degraded from the climax bluebunch wheatgrass-Sandberg's bluegrass steppe to a state of retrogression with dominance by cheatgrass or downy brome (the distinctive reddish or purplish colored straw). There was a lot (cover, density, biomass) of Sandberg's bluegrass along with forbs such as western yarrow and pale agoseris. Large patches of green vegetation in background were deeper microsites dominated by basin wildrye. The striking bone-white flowers in foreground were those of parsnip-flowered, or Wythe's wild-buckwheat. Snow wild-buckwheat was also present on this range (see below).

This was another "photo-quadrant" of vegetation on the same range shown in the immediately preceding two photographs.

Lincoln County, Washington. June (estival aspect) Dormancy and grain-shatter stage in Sandberg's bluegrass and cheatgrasss; peak standing crop and hard-dough stage in bluebunch wheatgrass; full-bloom stage in forbs. FRES No. 36 (Mountain Grasslands Ecosystem). K-44 (Wheatgrass-Bluegrass). SRM rangeland cover type designation was almost exclusively SRM 302 (Bluebunch Wheatgrass-Sandberg Bluegrass) which is the climatic climax Agropyron spicatum-Poa sandbergii (= P. secunda) zonal association of Franklin and Dryness (1973, p. 211) and also the the Agropyron spicatum-Poa secunda habitat type (Daubenmire, 1968, ps. iii, 18-20), which was in contrast to the lithosolic phase of this habitat type featured in the scablands range presented previously. This range was in some degree of degradation with much of the vegetation being a cheatgrass disclimax, the Bromus tectorum zootic climax of Daubenmire (1968, ps. 80-82). Local areas of the range shown here (eg. top of biscuit scab) were the edaphic climax Artemisia rigida-Poa secunda habitat type (Daubenmire, 1968, ps. iv, 40-42). Wheatgrass Series of or within Great Basin Shrub-Grassland biotic community or regional formation of Brown et al. (1998). Soils were primarily of the Anders-Bakeoven-Rock Outcrop Complex mapping unit; range sites for these two series were Loamy (L-2) and Very Shallow, respectively (Soil Conservation Service, 1981, ps. 11, 108, 110). The Loamy (L-2) range site (Anders series) predominated on the rangeland shown here. Columbia Plateau- Channeled Scablands Ecosystem, 10a (Environmental Protection Agency, undated).

106. An exclosure comparison- "Photo-quadrant" of vegetation just outside the channeled scablands range showed in the preceding three slides. Range vegetation shown in this slide was approximately ten to twelve feet away from the four-wire fence that enclosed the vegetation shown in the three preceding photographs. The rangeland vegetation shown in the current photograph was growing on an unmowed highway right-of-way such that it was vegetation protected from grazing (ie. the boundary or property line fence formed an exclosure that "fenced-out" rangeland which then served as reference vegetation). The conspicuous yellow composite was pale agoseris. The green grass was bluebunch wheatgrass and western or Columbia needlegrass. Dormant grass was Sandberg's bluegrass and cheatgrass. In this de facto exclosure cheatgrass was the least common grass species in contrast to its overall dominance in the disclimax of the pasture. Differences between range vegetation inside the enclosure (pasture) and that in the exclosure (adjoining highway right-of-way) was a textbook fenceline contrast.

Lincoln County, Washington. June (estival aspect) Dormancy and grain-shatter stage in Sandberg's bluegrass and cheatgrasss; peak standing crop and hard-dough stage in bluebunch wheatgrass; full-bloom stage in forbs. FRES No. 36 (Mountain Grasslands Ecosystem). K-44 (Wheatgrass-Bluegrass). SRM rangeland cover type designation was almost exclusively SRM 302 (Bluebunch Wheatgrass-Sandberg Bluegrass) which is the climatic climax Agropyron spicatum-Poa sandbergii (= P. secunda) zonal association of Franklin and Dryness (1973, p. 211) and also the the Agropyron spicatum-Poa secunda habitat type (Daubenmire, 1968, ps. iii, 18-20), which was in contrast to the lithosolic phase of this habitat type featured in the scablands range presented previously. This range was much closer to climax state or potential natural vegetation than that shown in immediately preceding three photographs. Wheatgrass Series of or within Great Basin Shrub-Grassland biotic community or regional formation of Brown et al. (1998). Soils were primarily of the Anders-Bakeoven-Rock Outcrop Complex mapping unit; range sites for these two series were Loamy (L-2) and Very Shallow, respectively (Soil conservation Service, 1981, ps. 11, 108, 110). Columbia Plateau- Channeled Scablands Ecosystem, 10a (Environmental Protection Agency, undated).

107. Pale agoseris, false dandelion, or mountain dandelion (Agoseris glauca var. glauca)- One of the plants of this species that was growing in the exclosure featured in the preceding photograph. This is just another of almost countless DYCs (Damn Yellow Composites) on North American range. Pale agoseris was one of the major forbs on this channeled scaplands range and was included to represent that grazing land. Pale agoseris is also one of the 200 species of range plants chosen for the International Intercollegiate Range Plants Contest. It was also shown here in that context. A good summary of A. glauca in context of Range Management was in the text/reference, North American Range Plants (Stubbendieck et al., 1992, p. 256-257).

Lincoln County, Washington. June (estival aspect); blooming and immediate post-bloom phenology

108. Silky lupine (Lupinus sericeus)- Many lupines call North American range their home. Silky lupine is one of the more widespread Lupinus species on the channeled scablands. Several views of this species were included at this juncture to present details of a characteristic channeled scablands range plant. The specimen introduced here was growing in a disclimax stand of cheatgrass, but also accompanied by cream, parsnip-flowered, or Wythe's wild-buckwheat and occasional plants of bluebunch wheatgrass and Columbia or western needlegrass.

This individual and its cheatgrass co-horts were growing on the depleted range described above

Lincoln County, Washington. June (late spring); full-bloom stage of phenology

109. Dreamy lupine and a creamy wild-buckwheat- Upper portion of shoot of the silky lupine plant in the preceding slide. The low shrub in full-bloom behind the full-bloom lupine was an individual of parsnip-flowered or Wythe's wild-buckwheat. Lincoln County, Washington. June (late spring); full-bloom stage of phenology.

110. Fruits and flowers on the scablands- Legumes and papilionaceous flowers produced on the silky lupine plant featured here. Lincoln County, Washington. June (late spring); full-bloom and immature fruit stages of phenology..

111. Snow wild-buckwheat (Eriogonum niveum)- Snow wild-buckwheat is a major--often dominant--low shrub on channel scablands range and other forms of Palouse Prairie steppe (see above also). Color of inflorescences in E. niveum varies as was demonstrated in this photograph. The larger specimen in the foreground had the bone-white color more common in this species while the smaller plant behind and to the right of the larger plant had blossoms of the pink (or pinkish) color phase. The Eriogonum species comprise one of the most widespread and species-rich genera of low shrubs in much of Intermountain North America. The wild-buckwheats are generally of very limited feed value, a notable fact given the common presence and often considerable cover on Intermountain ranges.

Lincoln County, Washington. June (late spring); full-bloom stage.

112. Look good enough to eat (almost)- The long flowering shoots and colorful inflorescences of snow wild-buckwheat. The pink (at least pinkish) tinge to petals of this species is due to peppermint stripe-like pigmentation. Presence of this colorful marking is a sure key to easy identification of this species (of course the off-white or bone-white coloration of petals in other plants of E. niveum complicate matters).

Lincoln County, Washington. June (late spring); full-bloom stage.

113. Parsnip-flowered, creamy, or Wythe's wild-buckwheat (Eriogonum heracleoides)- Another widespread.and, frequently, major low shrub on channeled scablands ranges is this species of wild-buckwheat. Parsnip-flowered initially but only superficially resembles snow wild-buckwheat with which it frequently grows beside on scab rangeland (as on the range shown in this photograph). Snow wild-buckwheat has much longer stems and less intricate flower clusters.

Lincoln County, Washington. June (late spring); full-bloom stage.

114. Flowers of parsnip-flowered wild-buckwheat- These two photographs presented detail and diagnostic features of the inflorescence of Eriogonum heracleoides. Comparisons of flower features shown in these two slides with those of E. niveum presented above reveal the distinct differences between these two often side-by-side channeled scablands shrubs.

Lincoln County, Washington. June (late spring); full-bloom stage

115. Sandberg's or Sandberg bluegrass (Poa sandbergii= P. secunda)- Several mature and now-dormant plants of Sandberg bluegrass growing on the first range presented above. These plants were part of this example of the Agropyron spicatum-Poa secunda habitat type, lithosolic phase (Daubenmire, 1968, ps. iii, 47-49). The more reddish (sometimes sort of purplish)-colored straw was cheatgrass. Rocks were basalt or similar igneous material.

Lincoln County, Washington. June (late spring); early dormancy, grain-shatter phenological stage

116. Sandberg's bluegrass and a more preferred composite- Immediately to the left of the conspicuous plant of Sandberg's bluegrass was a heavily grazed individual of narrowleaf skeleton weed or bush wire-lettuce. Species preferences and grazing selectivity of range animals was pronounced. Every individual plant of bush wire-lettuce observed by this author had a similar degree of use. It could not be determined if kind of range animal using narrowleaf skeleton weed was cattle, mule deer, rabbit, and/or some other species or combination of these, all of which had grazed this range. Bush wire-lettuce can grow into a large plant as was shown above when describing Palouse Prairie in the Loess Islands area. The comparative small size of plants on this shallow-soil site that had been grazed by cattle and wildlife with larger plants on the Loess Islands range that was grazed only by wildlife was obviously inconclusive (other than to show that some kind of range animal really liked bush wire-lettuce).

There was also a lot of cheatgrass on this range, much of the shoots of which had started to disintegrate. Most of the Sandberg bluegrass had received moderate degree of use. The photographer sought out ungrazed plants so as to present viewers with intack or complete plants for educational purposes as to species identification, habit of plant, size of inflorescence and sexual reproductive effort, etc.

Lincoln County, Washington. June (late spring). Early dormancy, grain-shatter phenological stage for Sandberg's bluegrass and pre-bloom stage for bush wire-lettuce (latter species had been maintained at a less mature stage by heavy grazing use).

117. Sandberg's bluegrass- Two individual plants of Sandberg's bluegrass in early dormancy and grain-shatter stage on channeled scablands range, Agropyron spicatum-Poa secunda habitat type, lithosolic phase (Daubenmire, 1968, ps. iii, 47-49). Reddish straw behind bluegrass plants was cheatgrass. The author included photographs of ungrazed Sandberg's bluegrass plants in order to portray morphology and features of this species. Such "selective grazing" by the photographer could foster the impression that Sandberg's bluegrass received less defoliation (lower degree of use defoliation) than cheatgrass, but this was not the case. Rather, cheatgrass had matured somewhat earlier (before) Sandberg's bluegrass and, as such, cheatgrass plants had disintegrated more than the later-maturing bluegrass.

Lincoln County, Washington. June (late spring).

118. Sandberg's panicles- Panicle inflorescences of Sandberg's bluegrass at or approaching grain-shatter stage on channeled scablands. Arrangement of spikelets on secondary branches of Sandberg's bluegrass were visible in the second of these slides. Lincoln County, Washington. June (late spring).

119. Bulbous bluegrass (Poa bulbosa)- This Eurasian annual or, frequently, short-lived perennial, grass is a widespread weedy species but it does not (other than at local scale) approach the impact (bad or good) of cheatgrass, the most important invader of this region. Bulbous bluegrass does not reach density, cover, biomass, etc. of cheatgrass and thus does not provide forage, soil cover, or fuel for wildfires as does the latter grass.

Latah County, Idaho. June (early summer); maturity and pre-bulb shatter stage.

120. Panicled bulbs- The common and scientific names of bulbous bluegrass indicate the unique reproductive feature of this annual grass. P. bulbosa produces one bulb or bulblet per spikelet even though there are characteristically three to eight florets per spikelet (Hitchcock and Cronquist, 1973, p. 658). Arnow (in Welsh et al., 1993, p. 856) gave a detailed and interesting account of this species, including its forage value, soil protection, and even use as companion crop in range seedings. Arnow also cited the finding that Poa bulbosa is the only grass to regenerate from true bulbs. This is one form of vivipary, but other grass species produce bulbils (Gould and Shaw, 1983, p.82).

121. Eye-catching panicle of a noxious species- Details of panicle of the Eurasian bulbous bluegrass. This was a close-up view showing spikelets in one of the panicles on the plant above the caption before last (two slide-set).

Niobrara County, Wyoming. Late June: grain shatter stage.phenology

Bulbous bluegrass also has shoots that have been described as "more or less bulbous at base" (Hitchcock and Chase, 1950, p. 122; Hitchcock and Cronquist, 1973, p. 658) so this morphological feature could be another or even the actual basis for names of this species. Most rangemen and foresters, however, focus on the most unique feature of bulb or bulblet production in the spikelet. Other grasses have an onion-like culm base.

Latah County, Idaho. June (early summer); maturity and pre-bulb shatter stage.

122. Namesake bulbs and bases- Rootcrowns on some plants of bulbous bluegrass illustrating basis of the specific epithet, bulbous, (and, thus also, common name) of this aggressive Eurasian grass. Niobrara County, Wyoming. Late June: peak standing crop of dead plants.

123. Thurber's needlegrass (Stipa thurberiana)- Thurber's needlegrass growing on the shallow-soil channeled scabland range at introduction of this section.This is one of the more important forage and, frequently, indicator species in drier parts of the Palouse Prairie Region, especially portions of the channeled scablands in the Columbia Plateau. Thurber's needlegrass is not a common species or even consistently found in channeled scablands plant communities, though does occur in this range vegetation as seen here.

Specimens in this and the next succeeding slide had immature panicles.

Lincoln County, Washington. June (late spring). Stage of phenology: hard-grain; immediate grain-shatter.

124. Inflorescences of Thurber's needlegrass- Examples of Thurbers needlegrass panicles on the above range that served as an example of climatic climax Agropyron spicatum-Poa sandbergii (= P. secunda) zonal association of Franklin and Dryness (1973, p. 211) which in this case was also the the Agropyron spicatum-Poa secunda habitat type, lithosolic phase (Daubenmire, 1968, ps. iii, 47-49).

Lincoln County, Washington. June (late spring). Phenological stage: hard-grain; immediate grain-shatter.

125. Shrub garland on a big scab- Around the talus sides of this relatively tall scab or small butte shrubs species syringa (Philadelphus lewisii) and western serviceberry (Amelanchier alnifolia) were thriving as what Franklin and Dryness (1973, p. 230) described as a shrub garland. This was talus slope vegetation had been described previously by Daubenmire (1968, p. 77). Just below ("downhill") the "rows" of these shrubs there were several plants of wax or squaw current (Ribes cereum) that also grew with various species of grass and forbs (presented below). The porous talus slopes and scattered piles of basalt and/or other igneous material function much like cistern so that this local habitat is more mesic than surrounding environments. This is basically the same ecolological phenomenon as that provided by lava flow malpais over various parts of the Western Range.

Shrub garlands are also similar to rock garlands, stripes, etc. in the alpine range type, sites, and ecosystems in form, structure, and general influences on plants and expressions of vegetation.

The distinctive geologic landmark shown here has been called variously a cinder cone or cinder butte. The series of Spokane floods that carved out the channeled scablands left this remnant of volcanic action (probably a lava flow).

More of these shrub garlands were presented below when discussing the channeled scablands land form in the Okanogan Drift Hills. Lincoln County, Washington. June (vernal aspect).

126. Syringa (Philadelphus lewisii)- Syringa or Lewis' mockorange growing on an igneous remnant that remained after the series of cataclysmic Spokane floods carried the rest of the lava flow away. Lincoln County, Washington. June (late spring). Full-bloom stage of phenology.

127. Forb-dominated range- Local area dominated by various forb at base of igneous talus on the side of a taller scab or small cinder butte. Cheatgrass, Sandberg's bluegrass, and bluebunch wheatgrass were also present. This was a local example of the herbacous vegetation presented in the two preceding photographs of a relatively large or tall scab (or small scablands butte). Forb species present included blanket flower, poker or roundleaf alumroot, and sticky geranium. A short distance away from this local plant community there was a small stand (perhaps one large clonal plant) of basin wildrye as shown in the first of the two photographs immediately above.

Lincoln County, Washington. June (late spring). Full-bloom phenological stage for most forbs.

128. Blanket flower (Gaillardia aristata)- This is a fairly common composite throughout much of the Columbia Basin. On channeled scablands it is most common on moister habitats. Lincoln County, Washington. June (late spring); full-bloom stage.

129. Poker or roundleaf alumroot (Heuchera cylindrica)- An attractive, interesting, and incidental range forb on channeled scablands range. This member of the Saxifragaceae also calls parts of the Great Basin home. Lincoln County, Washington. June (late spring).

130. Sticky geranium (Geranium viscosissimum)-There are a few Geranium species of economic and ecologtical importance across the Western Range. Most of these are of some forage value for livestock (depending on species) and probably of more utility as feed for wildlife, especially ruminants. Sticky geranium is one of the more common species on channeled scablands range. The individual presented in these photographs was growing in the local forb-dominated range plant community displayed above. Hermann (1966, p. 170) described this species as usually having a single stem whose herbage is "more or less sticky-glandular". He also reported that forage value of the geraniums "is evidently highly variable".

Lincoln County, Washington. June (late spring)

131. Wax or squaw current (Ribes cereum)- This shrub was also present in the greater or general range plant community associated with the tall scab or little butte with talus slopes described herein. The plant shown here was obviously loaded with fruit. This range shrub is a widely distributed across North America. It was selected as one of 200 species for inclusion in the Society for Range Management International Intercollegiate Range Plant Contest (Stubbendieck et al., 1992, ps. 424-425). (Incidentially, squaw current is more widely distributed and has a species range farther to the northwest, especially in Washington, than was shown in this publication.)

Browse value of wax current varies from poor for some kinds of livestock to good for some wildlife species (Stubbendieck et al., 1992, ps. 424). This species is well-adapted to the moist environment afforded by talus slopes. Examples of Ribes cereum growing on malpais range in the sagebrush shrub-steppe of eastern Oregon were presented in that chapter herein.

Lincoln County, Washington. June (late spring)

132. Channeled scablands marsh- Pothole lake supporting marsh vegetation in the complex of channeled scablands. The series of cataclysmic Spokane or Bretz floods had plunge pools where falling water created potholes or kolks of varying sizes, shapes, depths etc.(Allen et al, 1986, ps.23-28). These depressions or hollows that were scrated by the scouring, carving action of running flood waters over geologic time became natural lakes, ponds,or potholes. Kolks has also been used in reference to powerful underwater vortexes (giant whirlpools or malestroms) that scoured the potholes. It has been hypothesized that some of these vortices were formed when two flows of water moving in opposite directions and at two different speeds created powerful shear forces as they moved past each other. This is a different geologic process from the plunge pools that eroded away the columnar basalt parent material.

Some of these ponds or potholes formed by the Beretz floods were relatively large and deep while others were quite small and shallow. Thus persistence or permanence of water throughout the year varies widely and, of course, from year-to-year. Many of the smaller, shallower potholes or kolks pond water for brief periods and/or to such shallow depths that their range vegetation is either moist grasslands commonly dominated by basin wildrye, often with inland saltgrass, or as wetland plant communities. Some of these wetland range communities are ephemeral whereas others are have persistent surface water throughout the year (at least during the growing season) such that their range vegetation constitutes marshes of various species compositions, structures, and functions.

Several examples of these channeled scabland marshes were included variously throughout this section in association with adjacent larger and/or defining range sites, habitat types, and rangeland cover types. The marsh presented here was in the easternmost part of the channeled scablands at edge of the conifer zone in contrast to those presented below that were located to the west in the prairie zone (Yocom and Hansen, 1960, ps, 238-239).

Range vegetation of this marsh included two conspicuous wetland zones and one intermediate zone beween the wetland per se and adjoining bulebunch wheatgrass-Sandburg's bluegrass steppe. The zones of wetland plants were 1) spikerush, primarily common or creeping spikerush (Eleocharis macrostachya= E. palustris), in foreground and extending back in a v-shape pattern to water, and 2) hardstem, viscid, or great bulrush or, sometimes, tule used instead of bulrush (Scirpus acutus) closer to standing, ponded water (left and right of spikerush; at left and right margins of photograph). The zone between the bunchgrass prairie and the wetland or marsh vegetation proper was dominated by the Eurasian annual grass, medusahead wildrye. The inner-most edge of medusahead was in immediate foreground of slide. Background was varying landscape patches of bluebunch wheatgrass-Sandberg bluegrass steppe (some depleted to mostly stands of cheatgrass) and basin wildrye stands.

This marsh was in an ecotone between ponderosa pine (Pinus ponderosa) forest (more like woodland or even savannah) and bluebunch wheatgrass-Sandberg's bluegrass zonal association channeled scablands steppe. This range had been grazed by cattle a few weeks before this photograph was taken.

Spokane County, Washington. June (late spring). FRES No. 41 (Wet Grasslands Ecosystem). K-42 (Tule Marshes). No SRM. No appropriate unit given in Brown et al. (1998, p. 45), but it should be combination of Bulrush Series and Spikerush Series in a Columbia Plateau Intgerior Marshland biotic community. Columbia Plateau- Channeled Scablands Ecosystem, 10a (Environmental Protection Agency, undated).

133. Two zones in a marsh vegetation- Spikerush, primarily common or creeping spikerush, in foreground and great, hardstem, or viscid bulrush, to rear of spikerush, comprised two distinct zones of wetland range vegetation. This provided an example of zonal vegetation at local scale due to persistence of wet soil (tules or bulrushes being able to survive only with standing surface water remained longer into the growing season).

Spokane County, Washington. June (late spring). Peak-bloom to late anthesis phenological stage in bulrush.

134. Spikerush meets bulrush- Edge of range vegetation where the soil water-determined zones of great or viscid bulrush and creeping spikerush abutted each other on outer perimeter of a freshwater pothole marsh in an ecotone of ponderosa pine forest (or woodland or savannah) and bunchgrass prairie at eastern margin of channeled scaplands land form.

Spokane County, Washington. June (late spring)

135. Viscid or hardstem bulrush (Scirpus acutus)- Stand of hardstem, great, or viscid bulrush at peak standing crop and immediate post-anthesis phenological stage. In a freshwater marsh that developed on a pothole in channeled scablands adjacent to ponderosa pine-dominated forest. This marsh was in an ecotone of western yellow pine forest and bluebunch wheatgrass-Sandberg's bluegrass steppe.

Spokane County, Washington. June (late spring).

136. Inflorescences of hardstem bulrush- Two flower clusters of great or viscid bulrush in a pothole marsh at eastern perimeter of channeled scablands form of Palouse Prairie.

Spokane County, Washington. June (late spring)

137. Medusahead or medusahead wildrye (Elymus caput-medusae)- Another Eurasin annual weedy grass that naturalized in its adopted North America. These plants comprised a zone ("ring") at outermost (driest) edge of the hardstem bulrush-creeping spikerush dominated channeled scablands marsh. On an ecotone (transition zone) of western yellow (ponderosa) pine forest and bunchgrass prairie on which most common range vegetation was the zonal association of bluebunch wheatgrass-Sandberg's bluegrass.

Spokane County, Washington. June (late spring). Early bloom (pre-anthesis) stage.

138. Wetland in the scablands- Outer margin of a degraded bunchgrass prairie on which potential natural vegetation was a mosaic of the climatic climax bluebunch wheatgrass-Sandberg's bluegrass zonal association (Franklin and Dryness, 1973, p. 211) and the basin wildrye consociation (Clements, 1920, p.) interrupted by a freshwater tule or bulrush marsh in background.

The reddish-colored straw was a combination of cheatgrass or downy brome (dominant) and Sandberg's bluegrass (associate species) with isolated plants of scablands or stiff sagebrush. Large, green grass clumps were basin wildrye. It could not be determined what the climax or potential natural vegetation of the depleted range was as it appeared to be a combination of the edaphic climax Artemisia rigida-Poa secunda habitat type (Daubenmire, 1968, ps. iv, 40-42) and of a habitat type dominated by Elymus cinereus (Daubernmire, 1968). The pothole marsh was shown and described in more detail in the immediately succeeding slide.

Lincoln County, Washington. June (late spring); vernal aspect. FRES No. 41 (Wet Grasslands Ecosystem). K-42 (Tule Marshes). No SRM. No appropriate unit in Brown et al. (1998, p. 45) for the wetland, but there should be each of Bulrush Series, Rush Series, and Reed Canarygrass Series in a Columbia Plateau Intgerior Marshland biotic community (should be next to Great Basin Interior Marshland and Oregon I:nterior Marshland biotic communities). Likewise closest for the bluebunch wheatgrass-Sandberg's bluegrass steppe would Wheatgrass Series of or within Great Basin Shrub-Grassland biotic community or regional formation of Brown et al. (1998), but there should be a Bluegrass Series under that and some of this range vegetation would be of that designation. Better yet, there should be a Columbia Plateau Shrub-Grassland biotic community in Brown et al. (1998, p. 40). Habitat types given in previous paragarph. The marsh was Wet Meadow range site on a Cocolalla silt loam (Soil Conservation Service, 1981, ps. 22, 108). Range sites in foreground were a complex of Very Shallow and Loamy (L-2). Columbia Plateau- Channeled Scablands Ecosystem, 10a (Environmental Protection Agency, undated).

139. Summary view of a tule (bulrush)-Baltic rush-common spikerush pothole marsh- A freshwater marsh had developed on a depression deep and large enough to pond water throughout much of the warm growing season. Various edaphic-hydric zones of wetland vegetation comprised the range vegetation of this pothole or hollow marsh. This marsh differed from the preceding one in that the present wetland was in the prairie rather than at edge of ponderosa pine forest or savannah and eastern edge of channeled scablands.

This marsh closely fit the description of a mixed tule or bulrush marsh with American three-square or three-square bulrush (Scirpus americanus) joining great or viscid bulrush. Most importantly, this marsh included a large area (a circular zone) of Baltic rush (Juncus balticus) as well as a zone of creeing spikerush. At outermost edge of this diverse pothole marsh the introduced Eurasian, agronomic grass, reed canarygrass (Phlaris arundinacea) formed dense colonies. An example of this latter vegetational zone was in left foreground where it was partially fenced off from cattle (note height of reed canarygrass relative to height of studded steel T post).

Scabs or buttes of basaltic material formed a distant border around the marsh (background).

140. Feeding in a coulee- Not the Grand Coulee, but this wide draw (coulee) drained into the pothole marsh shown in the two preceding slides. Range vegetation in the first slide was the Elymus cinereus-Distichlis stricta habitat type of Daubenmire (1968, ps.iii,50-51) with the inland saltgrass having received relatively heavier degree of use than the basin wildrye.

Second slide featured a Sandberg's bluegrass-dominated area that had sizeable cover of cheatgrass and conspicuous western yarrow in foreground that gave way to the Elymus cinereus-Distichlis stricta; more spedifically, D. stricta var stricta) habitat type behind it. This second slide provided some scale to guage the size of basin wildrye when compared to a medium-sized, mature beef cow (predominately Angus breeding; approximatly 1200 pounds).

Lincoln County, Washington. June (late spring); vernal aspect. FRES No. 36 (Mountain Grasslands Ecosystem). K-44 (Wheatgrass-Bluegrass), but no K unit for basin wildrye. No SRM. For the Sandberg's bluegrass-dominated grassland in Brown et al. (1998, p. 40) closest was Wheatgrass Series of or within Great Basin Shrub-Grassland biotic community or regional formation), but there should be a Bluegrass Series under that or, better yet, there should be a Columbia Plateau Shrub Grassland biotic community with Bluegrass Series thereunder. Some of this range vegetation would be of that designation. Should also be a Basin Wildrye Series there also. Columbia Plateau- Channeled Scablands Ecosystem, 10a (Environmental Protection Agency, undated).

141. View from a pretty raw scab- On top of a rock outcrop on a typical scab various range sites and habitat types of channeled scablands range were visible and contrasts among these were striking. The strip of nearly bare land directly in front of the igneous rock cap and extending into far background was range dominated variously by Sandberg's bluegrass, cheatgrass, and Sanbdberg's bluegrass and scabland sagebrush. The latter was the edaphic climax Artemisia rigida-Poa secunda habitat type (Daubenmire, 1968, ps. iv, 40-42) whereas range dominated by cheatgrass was a cheatgrass disclimax or the Bromus tectorum zootic climax of Daubenmire (1968, ps. 80-82). Green patches were stands of basin wildrye. These wildrye populations were in swales or shallow hollows or small, local potholes that lost standing, surface water earlier in the growing season.

Whitman County, Washington. June (late spring). FRES No. 36 (Mountain Grasslands Ecosystem). Partially K-44 (Wheatgrass-Bluegrass). Partly SRM 106 (Bluegrass Scabland); no SRM for basin wildrye.For the Sandberg's bluegrass-dominated grassland in Brown et al. (1998, p. 40) closest was Wheatgrass Series of or within Great Basin Shrub-Grassland biotic community or regional formation), but there should be a Bluegrass Series under that or, better yet, there should be a Columbia Plateau Shrub Grassland biotic community with Bluegrass Series thereunder. Some of this range vegetation would be of that designation. Should also be a Basin Wildrye Series there also. A complex of soils and several range sites: primarily Bakeoven-Tucannon Complex soil mapping unit of Very Sahllow and Loamy (L-2) range sites respectively (Soil Conservation Service, 1980, ps. 82, 85) with local areas of Emdent or Pedigo silt loam comprising Alkali range site (Soil Conservation Service, 1980, ps. 30-31, 47-48, 82). Rock Outcrop mapping unit, front and center. Columbia Plateau- Channeled Scablands Ecosystem, 10a (Environmental Protection Agency, undated).

142. Pattern in the land; pattern in the vegetation (makes for a lot of pattern in the range)- The narrow, road- or corridorlike range site in the center of this slide was primarily of the edaphic climax Artemisia rigida-Poa secunda habitat type of Daubenmire (1968, ps. iv, 40-42). To either side of this narrow corridor on shallow soil was steppe vegetation dominated almost exclusively by basin wildrye (a basin wildrye consociation) or basin wildrye and inland saltgrass, the the Elymus cinereus-Distichlis stricta habitat type of Daubenmire (1968, ps.iii,50-51).

The extreme variation (or variety) in range sites and rangeland plant communities over short distances (small spatial scale) in the channeled scablands is some of the most extraordinary in North America. In this author's observation certain small riparian and the alpine ecosystems are about the only range types with greater variation in range at such small scale as the channeled scablands. Besides range sites, habitat types, and range cover subtypes there are numerous unique microsites or microhabitats often just steps from each other.

Whitman County, Washington. June (late spring). FRES No. 36 (Mountain Grasslands Ecosystem).Partially K-44 (Wheatgrass-Bluegrass). Partly SRM 106 (Bluegrass Scabland); no SRM for basin wildrye.For the Sandberg's bluegrass-dominated grassland in Brown et al. (1998, p. 40) closest was Wheatgrass Series of or within Great Basin Shrub-Grassland biotic community or regional formation), but there should be a Bluegrass Series under that or, better yet, there should be a Columbia Plateau Shrub Grassland biotic community with Bluegrass Series thereunder. Some of this range vegetation would be of that designation. Should also be a Basin Wildrye Series there also. A complex of soils and several range sites: primarily Bakeoven-Tucannon Complex soil mapping unit of Very Sahllow and Loamy (L-2) range sites respectively (Soil Conservation Service, 1980, ps. 82, 85) with local areas of Emdent or Pedigo silt loam comprising Alkali range site (Soil Conservation Service, 1980, ps. 30-31, 47-48, 82). Columbia Plateau- Channeled Scablands Ecosystem, 10a (Environmental Protection Agency, undated).

143.The deep and moist and the shallow and dry- On the example of channeled scablands shown here the foreground was a shallow hollow or pothole supported mesic range vegetation dominated by basin wildrye with inland saltgrass as associate species and spikerush (Eleocharis spp.), naturalized intermediate wheatgrass, and western yarrow (and, of course, some cheatgrass). This was the Elymus cinereus-Distichlis stricta habitat type (Daubenmire, 1968, ps.iii,50-51). Midground was the narrow, linear corridor-like range site(s) supporting alternatinv areas of cheatgrass disclimax, the Bromus tectorum zootic climax (Daubenmire, 1968, ps. 80-82) and the edaphic climax Artemisia rigida-Poa secunda habitat type (Daubenmire, 1968, ps. iv, 40-42). Behind this (right background) on deeper, more mesic soil was more the basin wildrye consociation and the basin wildrye-inland saltgrass. The pothole or depression in foreground was an Alkali range site on Emdent or Pedigo silt loam (Soil Conservation Service, 1980, ps. 30-31, 47-48, 82).

Whitman County, Washington. June (late spring). FRES No. 36 (Mountain Grasslands Ecosystem).Partially K-44 (Wheatgrass-Bluegrass). Partly SRM 106 (Bluegrass Scabland); no SRM for basin wildrye.For the Sandberg's bluegrass-dominated grassland in Brown et al. (1998, p. 40) closest was Wheatgrass Series of or within Great Basin Shrub-Grassland biotic community or regional formation), but there should be a Bluegrass Series under that or, better yet, there should be a Columbia Plateau Shrub Grassland biotic community with Bluegrass Series thereunder. Some of this range vegetation would be of that designation. Should also be a Basin Wildrye Series there also. Range site in foreground was Alkali on Emdent or Pedigo silt loam. Columbia Plateau- Channeled Scablands Ecosystem, 10a (Environmental Protection Agency, undated).

144. Bustin' through basin wildrye- Two examples of ungrazed basin wildrye (Elymus cinereus) showing the large size of this species in the channeled scablands. These plants were easily taller than a tall man's head (in a high-crowned hat). Elymu;s species are members of the barley or wheat tribe (Hordeae= Tritaceae) of subfamily Festucoideae.

Whitman County, Washington. June (late spring).

The next example of channeled scablands steppe range was in the level III ecoregion, Columbia Plateau- Okanogan Drift Hills (10d), in contrast to examples of scablands range in Channeled Scablands Ecoregion 10a that were described above. In general, range sites in the Okanogan Drift Hills area were noticably more mesic with deeper soils than were those in the Channeled Scablands Ecoregion. Obviously pothole marshes were hydric and basaltic rock outcrop was rock in both ecoregions and regardless of range site, habitat type, etc., but there were noticable differences.

145. Landscape-scale scene of more mesic, less varied channeled scablands steppe- The overall pattern of Columbia Plateau scabland range with its uplands and ridgetops, coulees and smaller draws, larger potholes with their marshes, and small malpais-like rock stripes was evident in these two "Big Picture" photographs in the Okanogan Drift Hills. This remarkable (and delightful) diversity in habitats that was due to differences in soils, general physiography, aspect, slope, and so forth with an array of range plant communities corresponding to range sites and habitat types was, however, less pronounced than in the more eastern channeled scablands of the Columbia Plateau that are closer to the Norther Rocky Mountains than to the Northern Cascades.

The "drift" feature of this land was obvious from the deep sand on hilltops. Also conspicuous was the landscape/landform feature of fewer scabs or small basalt buttes than in examples of channeled scablands discussed above. Otherwise the general scablands range type--complete with prominent erosion channels--was the same.

The two adjoining frames of this Okanogan Drift Hills range landscape included plant communities representative of the climatic climax Agropyron spicatum-Poa sandbergii (= P. secunda) zonal association of Franklin and Dryness (1973, p. 211) which included the Agropyron spicatum-Poa secunda habitat type (Daubenmire, 1968, ps. iii, 18-20) of bunchgrass prairie and also the climatic climax Artemisia tridentata/Agropyron spicatum zonal association of the sagebrush shrub steppe. There were scattered plants of basin wildrye (second slide) that appeared to coincide with more mesic microsites, most likely such microtopographic features as localized depressions or rock stripes, piles of basaltic rock, or other forms of lava flows that served as small wells much like malpais or talus slopes. There were also small very localized spots of cheatgrass disclimax, the Bromus tectorum zootic climax of Daubenmire (1968, ps. 80-82). An example of this along with (by side of) a local population of Sandberg's bluegrass was in foreground of second photograph (partially surrounding the large plant of basin wildrye). Western needlaegrass was widely scattered throughout this vegetation, but it was not a major species in any of the distinct range plant communities.

On the distant horizon or far background a bulrush-spikerush-Baltic rush marsh had developed around a large pothole. There were two pothole lakes in the distant background that were referred to below.

The showy low shrub in foreground of first photograph was the pink color phase of snow wild-buckwheat, a common shrub over much of the greater Palouse Prairie steppe and shrub-steppe region.

Douglas County, Washington. June (early summer), estival aspect. General or overall range ecosystem or landscape was FRES No. 36 (Mountain Grasslands Ecosystem). K-44 (Wheatgrass-Bluegrass), but there is no K unit for some of the local range plant communities. General rangeland cover type (Shiflet, 1994) was combination of SRM 302 (Bluebunch Wheatgrass-Sandberg Bluegrass) which is the climatic climax Agropyron spicatum-Poa sandbergii (= P. secunda) zonal association of Franklin and Dryness (1973, p. 211) and also the the Agropyron spicatum-Poa secunda habitat type (Daubenmire, 1968, ps. iii, 18-20) and of SRM 106 (Bluegrass Scabland). Brown et al. (1998, p. 40) did not provide a Columbia Plateau biotic community: closest vegetation unit was Wheatgrass Series of or within Great Basin Shrub-Grassland biotic community or regional formation (this would also require a Bluegrass Series under or for Great Basin Shrub-Grassland). Brown et al. (1998, p. 40) should have included a Columbia Plateau Shrub Grassland biotic community with Bluegrass Series, Wheatgrass Series, and Basin Wildrye Series thereunder. Columbia Plateau- Okanogan Drift Hills Ecosystem ,10d (Environmental Protection Agency, undated).

146. Another landscape-sized panarama of Okanogan Drift Hills steppe- The remarkable vegetational mosaic of channeled scablands range was evident in the panning view of the drift hills form of Columbia Plateau scabland range. Scabs were less common than on similar rangeland in the Channeled Scablands Ecosystem (Level III, 10a), but there was a similar channeled landform and many of the same species of range plants and range plant communities (similar range sites; most of same habitat types). Bunchgrass grassland in the foreground was representative of the bluebunch wheatgrass-Sandberg's bluegrass zonal association, a climatic climax (Franklin and Dryness, 1973, p. 211). The range plant community behind this (far right midground) was the edaphic climax and lithic association of Artemisia rigida/Poa sandbergii which coincides with or was interpreted as the Artemisia rigida-Poa secunda habitat type by Daubenmire (1968, ps. iv, 40-42).

Douglas County, Washington. June (early summer), estival aspect. General or overall range ecosystem or landscape was FRES No. 36 (Mountain Grasslands Ecosystem). K-44 (Wheatgrass-Bluegrass), but there is no K unit for some of the local range plant communities. General rangeland cover type (Shiflet, 1994) was combination of SRM 302 (Bluebunch Wheatgrass-Sandberg Bluegrass) which is the climatic climax Agropyron spicatum-Poa sandbergii (= P. secunda) zonal association of Franklin and Dryness (1973, p. 211) and also the the Agropyron spicatum-Poa secunda habitat type (Daubenmire, 1968, ps. iii, 18-20) and of SRM 106 (Bluegrass Scabland). There was also a representative range community of Artemisia rigida-Poa secunda habitat type (Daubenmire, 1968, ps. iv, 40-42) which was treated by the Society for Range Management (Shiflet, 1994, p. 6) as different--though often adjoining--SRM 106. Brown et al. (1998, p. 40) did not provide a Columbia Plateau biotic community: closest vegetation unit was Wheatgrass Series of or within Great Basin Shrub-Grassland biotic community or regional formation (this would also require a Bluegrass Series under or for Great Basin Shrub-Grassland). Brown et al. (1998, p. 40) should have included a Columbia Plateau Shrub Grassland biotic community with Bluegrass Series, Wheatgrass Series, and Basin Wildrye Series thereunder. Columbia Plateau- Okanogan Drift Hills Ecosystem ,10d (Environmental Protection Agency, undated).

147. Landscape view of channeled scablands in Okanogan Drift Hills- A patchwork of Palouse Prairie steppe range vegetation surrounded two pothole lakes in a more western portion of the channeled scablands. The range plant community in the rolling uplands shown in the foreground was a form of sagebrush-shrub steppe, the Agropyron spicatum-Poa secunda habitat type (Daubenmire, 1968, ps. iii, 18-20). Range vegetation in midground was a mosaic of local plant communities on microhabitats including basin wildrye consociations, bluebunch wheatgrass-Sandberg's bluegrass steppe, and sporadic "spots" of cheatgrass disclimax. There were individual plants of western needlegrass throughout this range vegetation, but they were nowhere near abundant and never dominant. Shrubs included big sagebrush, stiff or scablands sagebrush, rubber rabbitbrush, and both snow wild-buckwheat and parsnip-flowered, creamy, or Wythe's wild-buckwheat. The conspicuous range forb in foreground was some large individuals of silky lupine.

Douglas County, Washington. June (early summer), estival aspect. General or overall range ecosystem or landscape was FRES No. 36 (Mountain Grasslands Ecosystem). K-44 (Wheatgrass-Bluegrass). General rangeland cover type (Shiflet, 1994) was combination of SRM 302 (Bluebunch Wheatgrass-Sandberg Bluegrass) which is the climatic climax Agropyron spicatum-Poa sandbergii (= P. secunda) zonal association of Franklin and Dryness (1973, p. 211) and also the the Agropyron spicatum-Poa secunda habitat type (Daubenmire, 1968, ps. iii, 18-20) and of SRM 106 (Bluegrass Scabland). Also as just featured the Agropyron spicatum-Poa secunda habitat type (Daubenmire, 1968, ps. iii, 18-20). Brown et al. (1998, p. 40) did not provide a Columbia Plateau biotic community: closest vegetation unit was Wheatgrass Series of or within Great Basin Shrub-Grassland biotic community or regional formation (this would also require a Bluegrass Series under or for Great Basin Shrub-Grassland). Brown et al. (1998, p. 40) should have included a Columbia Plateau Shrub Grassland biotic community with Bluegrass Series, Wheatgrass Series, and Basin Wildrye Series thereunder. Columbia Plateau- Okanogan Drift Hills Ecosystem ,10d (Environmental Protection Agency, undated).

148. Plentious pothole- A large pothole (depression, hollow, swale, or other term) carved millenia ago by the great cataclysmic Spokane or Bretz floods was now home to this beautiful and very productive Columbia Plateau grassland. Basin wildrye and inland saltgrass combined to develop into this mesic expression of Palouse Prairie. Frequently basin wildrye forms single-species stands (expansive populations or colonies), but on other range sites this large--often immense--member of the barley tribe (Hordeae= Tritaceae) "teams up" with the low-growing eragrostoid saltgrass, member of the small tribe, Aeluropodeae, to form this interesting two-layers-of-grass grassland.

Clements (1920, p. 151) viewed Elymus condensatus (= E. cinereus) as "a frequent associate" except on saline lowlands where it had to be regarded as "a subclimax dominant". In Clementsian monoclimax theory the terminus of vegetation on the sere that was typical of mesophytic midslopes across a vast space of land--a region--was the final vegetational product (plant community) due to prevailing climate once a peneplain was formed when hilltops were leveled and lowlands filled over course of geologic time. At that point in the geologic cycle the climatic (= regional or zonal) climax would prevail. Current climaxes on floodplains, moist mountains, and other especially favorable habitats were subclimax to the final climatic climax across the region. In the Palouse Prairie, basin wildrye would be subclimax to bluebunch wheatgrass, the major dominant throughout the Palouse Region (Clements, 1920, ps. 149-150). In polyclimax theory, basin wildrye forms an edaphic climax. Elymus cinereus-Distichlis stricta constitute a soil association in contrast to zonal associations that are climatic climaxes (Franklin and Dryness, 1973, ps. 211, 226).

Douglas County, Washington. June (early summer), estival aspect. Primarily soft- to hard-dough stage for both grass species. FRES No. 36 (Mountain Grasslands Ecosystem). No Kuchler unit for this climax grassland. No SRM either. Nor did Brown et al. (1998) provide a classifiction unit. Brown et al. (1998, p. 40) should have included a Columbia Plateau Shrub Grassland biotic community with a Basin Wildrye Series thereunder. What gives folks? Daubenmire (1968, ps. iii, 50-51) rode to the rescue with the Elymus cinereus-Distichlis stricta habitat type. Columbia Plateau- Okanogan Drift Hills Ecosystem ,10d (Environmental Protection Agency, undated).

149. Great natural pasture and great biodiversity in the scablands- In this floor of a channel and gradually progressing up to a ridgetop of lava rock stripes in the western channeled scablands numerous plant species constituted a diverse range community that provided a "smorgasboard" of forage and browse. Important and indicative species of grasses included bluebunch wheatgrass, Sandberg's bluegrass, western or Nelson's needlegrass (Stipa occidentalis var. nelsonii= S. nelsonii), squirreltail bottlebrush, and, as always, cheatgrass. Big sagebrush and rubber rabbitbrush (along with snow wild-buckwheat and parsnip-flowered, creamy, or Wythe's wild-buckwheat) were widely scattered among the obviously more abundant (cover, density, frequency, etc. parameters) grasses, the dominant of which was bluebunch wheatgrass. This was clearly grassland and not a shrub-grass savanna (ie. steppe not shrub-steppe vegetation). The most common--though it was quite uncommon--forb in the channel was western yarrow.

The range vegetation up on the sides and top of ridges was different from that of the channel floor, but while these were two range sites both likely fell into the same rangeland cover type. Details of ridge vegetation were given in the next set of two slides.

Douglas County, Washington. June (early summer), estival aspect. Peak standing crop. General range ecosystem or cover type was FRES No. 36 (Mountain Grasslands Ecosystem). K-44 (Wheatgrass-Bluegrass). The Society for Range Management rangeland cover type that most closely fit this range plant community appeared to be SRM 101 (Bluebunch Wheatgrass). Did not readily fit into any Daubenmire (1968) habitat type without streatching to a bluebunch wheatgrass -big sagebrush shrub steppe which this was not. Columbia Plateau- Okanogan Drift Hills Ecosystem ,10d (Environmental Protection Agency, undated).

150. Flowers aplenty but the garland was one of shrubs- The first of these two photographs showed some of the area on which the range plant community of the channel floor met that of the ridge. If this was an ecotone it was one of abrupt edges. The foreground was of the bluebunch wheatgrass-dominated steppe that was shown and described immediately above. Here at the upper part of the channel (higher elevation; farthest up the slope) cover and density of big sagebrush and rubber rabbitbrush were substantially greater than in the channel floor. On the ridge, especially the top thereof, both bluebunch wheatgrass and big sagebrush were less abundant than in the channel. A major--clearly an associate--species up on the ridge was arrowleaf balsamroot (Balasmorhizza sagittata). The tall shrub was western serviceberry.

The western serviceberry-arrowleaf balsamroot ridge community had developed along a basalt rock stripe. Consistent with Bertz' view of channeled scablands as "wounds only partially healed" (Allen et al., 1986, p. 31) range vegetation and this low outcropping of igneous rock could be understood as a "better-healed wound" or a "wound that was closer to being scabbed over". This plant community was a form of shrub garland (Daubenmire, 1968, p. 77; Franklin and Dryness, 1973, p. 230) similar to but less species-rich than the one presented previously. Daubenmire (1968, p. 77) explained that stands of western serviceberrry in the Artemisia-Agropyron zone area are "a distinctive talus vegetation" due to rapid penetration and efficient storage of precipitation with subseqwuent reduced evaporation. These elongated and narrow habitats with their porous, rocky substrata support range vegetation that is more mesophytic than that of adjacent rangeland, hence a garland of shrubs in an expanse of grassland having scattered smaller species of more xeric shrubs.

Rock stripes function much like natural cisterns in the same fashion as malpais in areas of previous volcanic activity.

Douglas County, Washington. June (early summer), estival aspect. Peak standing crop. There were not appropriate FRES, Kuchler, or SRM units for this unique and limited range vegetation (ie. too small to be covered by major descriptive and map units of potential natural vegetation).This edaphic climax vegetation was a distinctive part of channeled scablands range. Columbia Plateau- Okanogan Drift Hills Ecosystem ,10d (Environmental Protection Agency, undated).

151. Scablands chums- Two photographs of two co-dominant grasses of channeled scablands range. On some range sites and habitat types of the channeled scablands one-spike or single-spike oatgrass (Danthonia unispicata) joins Sandberg's bluegrass for co-dominance of climax range vegetation. These two slides introduced these cespitose species as co-dominants of the channeled scablands form of Palouse Prairie. In the first ((horizontal) of these two slides single-spike oatgrass was on the left and Sandberg's bluegrass was on the right. Position of these co-dominants was reversed in the second (vertical) slide: Sandberg's bluegrass at left, single-spike oatgrass at right and slightly to rear.

Sandberg's bluegrass is one of the major, most widely distributed grasses of the channeled scablands typically being a dominant, co-dominant, of widespread associate on scablands ranges. Sandberg's bluegrass is commonly a decreaser being an important--again, often a dominant--species of steppe vegetation depending on range site, habitat type or range type. On some range sites of Palouse Prairie grasslands Sandberg's bluegrass is an increaser and high cover and density of Poa sandbergii (P. secunda) indicates disturbance (eg. drought) or range abuse (eg. overgrazing).

On some range sites single-spike oatgrass is a side-by-side co-dominant to Sandberg's bluegrass (as suggested by these two photographs). On other ranges--depending on range site and grazing history--Danthonia unispicata forms consociations and occurs as single-species stands (more like expansive populations) or nearly so with occasional plants of cheatgrass and/or Sandberg'sbluegrass. An example of the latter species composition and structure was presented in the immediately following slide. Single-spike oatgrass is not nearly as common or frequent a dominant as is Sandberg's bluegrass as indicated by the Society for Range Management (Shiflet, 1994, p.6 ) rangeland cover type designation (and description) of Bluegrass Scabland (SRM 106). Stated another way: one-spike oatgrass is a distant second among equals on bluegrass scabland range, other than locally (local scale) on some ranges.

Spokane County, Washington. June (early; summer); hard-grain, immediate pre-shatter phenological stage.

152. Single-spike oatgrass consociation- Example of a sizeable population of Danthonia unispicata on a channeled scablands range. Few--precious few--plants of Sandberg's bluegrass and cheatgrass grew in what was otherwise a population of one-spike oatgrass. Rangeland plant communities like this (consociations of Danthonia unispicata) are relatively scarce throughout the channeled scablands of the Palouse Prairie. At least such stands are not nearly as widespread as those dominated by Sandberg's bluegrass or co-dominated by bluebunch wheatgrass and Sandberg's bluegrass.

In a similar vein relative to species' importance it bore noting that across North American range Danthonia unispicata is not as important a species as are D. californica, D. intermedia, or D. parryi which are the three oatgrass species included on the Society for Range Management International Intercollegiate Range Plant Contest. Nonetheless, one-spike oatgrass is an important and, locally, a major grass on channeled scabland steppe range. Single-spike oatgrass is clearly a defining species of the channeled scablands form of Palouse Prairie.

Spokane County, Washington. June (early; summer); hard-grain, immediate pre-shatter phenological stage. FRES No. 36 (Mountain Grasslands Ecosystem). K-44 (Wheatgrass-Bluegrass). SRM 106 (Bluegrass Scablands). There was not an appropriate classification unit of this range vegetation in Brown et al. (1998). Columbia Plateau- Channeled Scablands Ecosystem, 10a (Environmental Protection Agency, undated).

153. Plants of single-spike or one-spike oatgrass (Danthonia unispicata)- General appearance and morphology of this cespitose grass were shown in the plants presented in these two photographs.

Spokane County, Washington. June (early; summer). Plants were at full maturity going into dormancy having completed this year's life cycle. Sexual state of phenology: hard-grain, immediate pre-shatter.

154. Features of one-spike oatgrass- Overall structure and arrangement of shoots of single-spike oatgrass were presented in the these two slides, including panicle inflorescences on numerous sexual shoots.

155. Shoots of one-spike oatgrass- These two photographs displayed more details of leaf and culm including the interesting pilose pubescence on internodes that characteristically covers much of the leaves also.

Spokane County, Washington. June (learly; summer). Plants were at full maturity going into dormancy having completed this year's life cycle. Sexual state of phenology: hard-grain, immediate pre-shatter.

156. Spikelets of one-spike oatgrass- Common and scientific names of this species refer to the characteristic single spikelet in its panicle inflorescence. Less commonly there are two or even three spikelets. When more than the single terminal spikelet are pressent, the lower spikelet(s) are appresseed (ie. hug the culm) in contrast to the typically elongated pedicel of the spikelet at terminus of sexual shoots (Hitchcock and Chase, 1950, p. 312).

The Danthonia species in North America produce cleistogamous(self-fertilized, unopened) spikelets in their lower sheaths (Hicthcock and Chase, 1950, p. 307). This photographer could not find good examples of cleistogamy in the D. unispicata specimens growing on this range.

157. Just what the Palouse Prairie needed: another alien, annual bromegrass- Hairy brome or hairy chess (Bromus commutatus) forming a dense stand that is typical of this Eurasian annual species. Hairy chess is just one of several such annual invaders that made it to North America complements of the white man and, thanks to his continued disturbances, one that continues to be successful (though minor) weed across much of the Palouse Prairie. Fortunately this annual brome has been a much less successful invader than downy brome (cheatgrass). This local stand of B. commutatus was in the Palouse Hills of the Northern Rocky Mountains portion of the Palouse Region.

Latah County, Idaho (June, early summer). Phenology: full maturity, dead, grain-shatter stage.

158. Columbia Plateau common reed-saltgrass marsh- Some of the potholes and subsequently formed lakes created by the series of Spokane or Bretz cataclysmic floods resulted in development of marshes dominated by common reed (Phragmites communis= P. australis) as the upper herbaceous layer of range vegetation. The lower herbaceous layer (of vascular range plants) in reed marshes is usually inland saltgrass (the same as in basin wildryed-dominated mesic grasslands). On some parts of this freshwater marsh inland saltgrass was more of an associate species to the much larger common reed. Alternatively, saltgrass could be interpreted as dominant of the lower herbaaceous layer of this range wetland. On the outer edges or periphrey of this marsh, inland saltgrass formed a single-species sward. Apparently this zone of range vegetation was beyond the boundary of or out of the marsh per se. It could be argued that such zones of saltgrass were, precisely speaking, out of the wetland and instead constituted an ecotone or transition from wetland to steppe vegetation.

Common reed is probably the most widely distributed angiosperm on Earth. This distribution, however, is a discontinuous or disjunct species range; more specifically, a diffuse species range (or range pattern). Common reed occurs on every continent (except Antarctica) and its use by humans traces to some of the earliest prehistory. In recent years it was discovered that non-native genotypes of P. communis= P. australis had apparently naturalized in parts of North America with the result that these exotic genotypes had become major, aggressive weeds. Genomic studies indicated that there are now both native and non-native (introduced from Eurasia) haplotypes of common reed. Apparently all common reed genotypes reproduce as clones as well as also producing viable seeds so that reproduction is (can be) both sexual and asexual.

Non-native common reed generally produces larger individual plants (shoots) and denser stands. This genetic form of the species appears to be an aggressive weed that threatens native plants. Hence the spread of non-native P.communis has become a matter of concern for natural areas, national parks, and wildlife refuges. It could become a major weed in regard to other lands and land uses. The state of Washington proposed listing and management of non-native genotypes of Phragmites as a Claqss C weed (Washington State Noxious Weed Control Board, 2003).

Most or all of the P. communis shown in these and the following photograph was probably a halotype that is not native to North America. However, it certainly was the same species as the native common reed. Thus, range vegetation shown here was representative of the native common reed-inland saltgrass marsh vegetation that is indigenous to the channeled scablands. Marshes have received relatively little coverage in the native vegetation literature of the Pacific Northwest, including that for Oregon and Washington. As such there is not much available material in regard to reed-dominated (or even bulrush-dominated) marshes within the greater Palouse Prairie Region.

Historic treatment of "reed swamps" by pioneer plant ecologists like F.E. Clements and A. Tansley was discussed elsewhere in this publication under both literature review of development of the range type concept and treatment of the common reed communities along the Rio Grande (Miscellaneous Grasslands chapter).

Most of the shrubs at far margin of this freshwater marsh were the exotic and dreadfully noxious saltcedar (Tamarix parviflora according to Hitchcock and Cronquist, 1973, p.296; perhaps T. ramosissima is the more nearly correct species designation). Control and ideally--though not realistically--eradication of this brush species would eeem as much if not more desirable and pressing than control of non-native genotypes of a native grass.

Grant County, Washington (June (early summer). Pre-bloom stage of common reed; panicles and dead shoots were "left over" from previous previous year(s). FRES No. 41 (Wet Grasslands Ecosystem). No Kuchler unit. No SRM. No Columbia Plateau Interior Marshland in Brown et al. (1998, p. 45). Columbia Plateau- Yakima Folds Ecoregion, 10g (Environmental Protection Agency, undated).

159. Common reed bed- Phragmites communis= P. australis dominating a freshwater channeled scablands marsh. Morphological features such as thick shoots and the dense nature of stands suggested that this was more likely than not to be a halotype that is not native to North America. This cannot be definitively determined on the range.

Grant County, Washington (June (early summer). Pre-bloom stage; panicles and dead shoots were those of previous year(s).

160. Inland saltgrass (Distichlis stricta, specifically D. stricta var stricta )- One characteristic plant of inland saltgrass growing as co-dominant to common reed on a channeled scablands pothole marsh in the Yakima Folds Ecoregion portion of the Columbia Plateau. Grant County, Washington (June (early summer). Grain-set phenological stage.

Basin or giant wildrye (Elymus cinereus) range type

All human organization of things is imperfect and as such has misfits. Basin wildrye was a misfit in this publication on range types. Giant wildrye simply did not fit readily or logically anyplace. This species was placed where it seemed to be the least out of order. No, it still ain't in order, but it is in the publication.

Basin wildrye does occur in the sagebrush-shrub steppe as for instance along watercourses. This species also grows in the salt desert shrub of the Great Basin Desert (Barbour and Billings, 1988, ps. 220-221). Young and Sparks (2002, p. 51) referred to "extensive areas" of Great Basin wildrye in moist and/or saline sites in the Great Basin. Kagan et al. (2004) with the Oregon Natural Heritage program recognized two Great Basin wildrye plant associations: 1) basin wildrye bottomlands (Leymus {Elymus] cinereus) and 2) giant wildrye alkali marsh- saltgrass (Leymus [Elymus] cinereus-Distichlis spicata). Basin wildrye also grows along streams and in moist openings in the western yellow pine (Pinus ponderosa) forest. Giant wildrye is also a local range type in the channeled scablands of the greater Palouse Prairie. Giant wildrye was a major range community only in the channeled scablands and not in any of the other larger vegetation units.

Basin wildrye stands (consociations and mixed-species communities) are grasslands, small in area though the remaining relicts are. Therefore, the little spatial-scale communities of giant wildrye in the High Lava Plains and Blue Mountains provinces (Franklin and Dyrness, p. 6) of Oregon were place in the Palouse Prairie and Miscellaneous "chapters". This natural single-species range grass community was placed herein under Grasslands consistent with organization of this publiction as to biomes at the broadest community level.

Basin wildrye communities in channeled scablands range were covered previously (above) that subheading in this chapter devoted to the Palouse Prairie. While this spatial arrangement of basin wildrye appeared disjunct it permitted placement of range plant communities that are dominated by this valuable, high-yielding, native grass withih geographic units. This organization presented occurrence of basin wildrye grasslands within their larger or more general zonal range vegetation and in spatial proximity to adjacent range types.

161. "Shades of the Past": consociation of Great Basin, basin, or giant wildrye (Elymus cinereus). This relict stand of basin wildrye was but a remnant example (relatively large though it was) of what was once a major climax plant community in much of the Intermountain Range Region. Grassland comprised of natural single-species stands of basin wildrye or of basin wildrye and inland saltgrass (Distichlis stricta) was often the most productive natural (pre-Columbian) vegetation of the Great Basin and parts of adjoining areas. This remarkable high yield was in terms of both biomass (= herbage) and useful forage. In frontier times basin wildrye was cut for hay and was the largest source of harvested forage for U.S. Army cavalry horses (A.L. Lesperance, personal communication). Young and Sparks (2002, p. 51) mentioned "extensive areas" of Great Basin wildrye dispersed with wetlands, salt desert scrub, and other Basin and Range vegetation.

In Classification of Native Vegetation of Oregon by the Oregon Natural Heritage group (Kagan et al., 2004) two Great Basin wildrye plant associations were recognized: 1) basin wildrye bottomlands (Leymus [= Elymus] cinereus) and 2) giant wildrye alkali marsh-saltgrass (Leymus [= Elymus] cinereus- Distichlis spicata). The example of giant wildrye shown here was the basin wildrye bottomlands association.

Undoubtedly much of the original climax basin wildrye-dominated vegetation was destroyed to make way of farming and other land uses. Much of the remaining giant wildrye range was overgrazed and/or overmowed for hay. As a result relatively little of the original basin wildrye range vegetation remains.

The relict vegetation shown above was in the Harney Basin of the High Lava Plains province (Franklin and Dyrness, ps. 6, 32-34). This extensive stand was one of the few tracts of its species that remained in the former grassland-marsh complex that developed in the Silvies River valley. Inland saltgrass was absent from this stand which again was an example of the basin wildrye bottomlands form of this type.

Efforts continue to restore this basin wildrye to some of the land once dominated by it. Magnar and Trailhead are two accessions of Great Basin wildrye that were selected and released by Soil Conservation Service Plant Materials Centers. Seed is available from several seedsmen.

Harney County, Oregon. June. Mid-bloom phenological stage. Basin wildrye bottomland association of Kangan et al. (2004).Northern Basin and Range- High Desert Wetlands Ecoregion, 80e (Thorson et al., 2003).

162. Giant wildrye in the woods- Basin wildrye is widely distributed in northwestern North America. In its geographic range basin wildrye grows on diverse habitats varying from Northern Great Plains grasslands southward into wet and fairly saline sites in Great Basin vegetation. Between these latitudinal extremes giant wildrye grows in mesic environments in mountain forests. Shown here was an example of a consociation of basin wildrye had developed on a moist yet well-drained opening within a ponderosa pine (Pinus ponderosa) forest in the Blue Mountains province (Franklin and Dyrness, 1973, ps.6, 27-29). Based on the Orgeon Natural Heritage Classification of Native vegetation of Oregon (Kagan et al., 2004) this was a basin wildrye bottomlands plant association, there being no inland saltgrass in this specific community and no vegetation unit for this basin wildrye in ponderosa pine forest.

Grant County, Oregon. June. Soft-dough phenological stage. Variant of basin wildrye association of Kagan et al. (2004). Blue Mountains-John Day/Clarno Uplands Ecoregion, 11a (Thorson et al., 2003).

Northern Mixed Prairie-Palouse Prairie Ecotone or Transition Grassland

The mixed prairie of the Northern Great Plains and the Palouse Pairie of the Northern Rocky Mountain Region come into contact in northern, especially northeastern, Wyoming and from southeastern Montana on northwest-ward (Weaver and Albertson, 1956, p. 340). Therein these authors quoted Clements (1920) as describing how "the bunch-grass prairie passes so gradually into the mixed prairie in central Montanan that no line can be drawn between them". The transition is from dominance by needle-and-thread and western wheatgrass to bluebunch wheatgrass. Weaver and Albertson (1956, p. 340) elaborated further by citing the work of Heady (1950) which showed that optimum development of the Palouse Prairie in Montana was in foothills between the Rocky Mountains and Great Plains.

The short treatment that followed dealt with this ecotonal phenomenon. As this ecotonal or transition grassland could fit both Palouse Prairie and Northern Great Plains Grassland (or be unique and distinct from both) this section was included in both the chapters on Palouse Prairie and Northern Great Plains.

163. Hybrid Grassland- A form of transition or ecotonal grassland between Northern Great Plains mixed prairie and Palouse Prairie. The potential natural vegetation was a mixture of Palouse Prairie co-dominants, bluebunch wheatgrass and Idaho fescue, and mixed prairie dominants, western wheatgrass, green needlegrass, and needle-and-thread.

This particular range had been greatly modified by human action and in addition to the native grass species just listed, smooth bromegrass and crested wheatgrass (introduced Eurasian perennials that are naturalized throughout this region) had become part of "the mixture".

The major forb on this grassland, which was home to pronghorn as well as cattle, was silver (or silvery) or tailcup lupine (Lupinus argenteus var. argenteus).

Sheridan County, Wyoming. Mid-June; flowering stage for bulebunch wheatgrass and Idaho fescue. Vegetational classification of this highly human-modified grassland would be meaningless. Northwestern Great Plains- Mesic Dissected Plains Ecoregion 43q (Chapman et al., 2004).

Restoration of Palouse Prairie

It was noted above that the eastern forms of Palouse Prairie--especially that on arable, often highly erodible, land-- such as that remaining in the Northern Rocky Mountains is regarded as one of the most endangered and most drastically reduced natural ecosystems in the United States (Johnson and O'Neil, 2001, p. 49). Sadly for rangemen, wildlifers, and native plant enthusiasts, this is an example of the ironclad Law of Succession Land Uses: land moves to its level of highest economic return (the land use that generates greatest profit per area of land). In the Palouse Prairie soil that formed over millions of years from bunchgrass prairie was far more valuable in terms of agricultural economic return than the pasture afforded by the native steppe or shrub-steppe vegetation. Unfortunately, the Law of Succession of Land Uses was violated all too often when human passions (including greed and irrational optimism) overrode pecuniary reality resulting in economic and ecological disaster when Palouse Prairie on marginal, erosion-prone land was converted into cropland.

In critical periods throughout human history misuse of the plow resulted in financial ruin and depletion of natural resources along with collapse of man's social order. At certain junctures in the course of man, well-to-do so nations with a sense of destiny and a feeling of obligation to their grandchildren (or when remedial action became unquestionably essential) mustered the moxey to save what was left of their once abundant natural heritage. Periodically, people made the necessary sacrifices for the cause of conservation. Sometimes this resulted in reinvestments in grassland, wetland, and forest restoration.

In the United States the Conservation Reserve Program (CRP) has been one of the most successful of these reinvestments in God-given natural wealth. The CRP is not perfect (far from it). It does subsidize undue and misplaced optimism and to a large degree it rewards poor stewartship. The CRP remains, however, the best approach that is politically and economically acceptable by which marginal farmland has been retired and restored to grassland, wetlands, forests, or some form of permanent vegetation that protects soil and water, the most basic natural resources of agriculture. Some--not near enough--of former Palouse Prairie has been reseeded to native grasses resulting in grassland that approaches (more-or-less) the virgin vegetation of the bunchgrass prairie. Two examples were presented here.

164. Restoration of the mesic regional dominant of the Palouse Prairie- Single-species stand of Idaho fescue on former cropland in the Palouse Rrairie Region. Make-up of plant species reseeded on this field was not known to this author, but the only species present on the land at time of photograph was Idaho fescue. It could not be determined if this was the only species planted or if a mixture had been seeded. Mixtures are almost always superior single-species seedings so it was a reasonably safe assumption that several species had been recommended for this reseeding project. Perhaps several plant species were drilled and Idaho fescue was the only one that emerged or persisted.

Even a monoculture of one of the two regional dominant species (bluebunch wheatgrass being the other of course) constituted partial restoration of Palouse Prairie grassland. Likewise, age of this stand was unknown such that it might be possible that other species were seeded and had yet to emerge and become established. In matters of range reseedings rangemen can compromise and "settle for half a loaf". A re-established stand of the more mesic of two dominants of the climatic climax is plenty cause for celebration. All the more given that this was such a fine dense stand.

It should be underscored that Idaho fescue commonly existed as a consociation and resembled a "natural monoculture" on the virgin Palouse Prairie. Photographs of two such one-species steppe communities were presented above, including one from the same county in which this grand stand was thriving.

Whitman County, Washington. June.

165. Takes a close look to tell- A grain field in the Palouse Prairie Region that was reseeded under terms of the Conservation Reserve Program. The resulting restored grassland was a mixture of Idaho fescue and bluebunch wheatgrass, the two regional dominant species of the Palouse Prairie. Composition of the seeding mixture that was planted was not known to this photographer. There were a few widely scattered plants of intermediate wheatgrass. These were more common around edges of the field suggesting that these intermediate wheatgrass plants were adventive or only able to establish or persist where competition with the native dominants was less intense. Whatever the mixture, seeding practices, and other management measures this was an amazing accomplishment in reclamation of bunchgrass prairie.

The characteristic spatial distribution of native cespitose grasses on Palouse Prairie steppe usually presents itself as a regular pattern of dispersion. This roughly equi-distant dispersion (much like trees in an orchard, corn planted in hills, crosses in a military graveyard) suggest or brings to the imagination the appearance of "rows", as in this case of bunchgrasses. Thus, even a natural steppe in the Palouse Prairie resembles crops planted in rows or hills. When looking for natural steppes rangemen pay most attention to species composition (along with structure and physiogonomy) of the vegetation. Given the relatively simple species make-up of native steppe (and of habitat types thereof), such as natural single-species stands, the student of the range must look closely at plant spacings when determining naturalness of grasslands. This reseeded grassland was an anthropogenic restoration of the Agropyron spicatum-Festuca idahoensis climatic climax zonal association (Franklin and Dryness, 1973, p. 211). The presence of grass plants in drill rows was obvious in the example shown here. Individual grass plants where closer together in one direction than in others. This was the direction in which the drill was pulled behind the tractor (ie. directional orientation of drill rows).The observant, knowledgable rangeman would determine quickly that this was a range reseeding project (and a completely successful one). Familarity with agricultural programs would (did) lead one to strongly suspect a Conserve Reserve Program reseeding. Consultation with folks at the district office of the Natural Resources (Soil) Conservation Service confirmed the suspicion.

Absence of native perennial forbs such as western yarrow was another indicator that the grassland presented here was the result of artificial planting and not natural regeneration via secondary plant succession. This was not "cut-and-dried". The large, green, clumped forbs were bush wire-lettuce which are native and common throughout much of the Palouse Prairie.

This tract and the reseeded land of the Palouse Prairie shown in the immediately precceding photograph were "picture postcard-perfect" examples of the Conservation Reserve Program in action. Grassland restoration. Reclamation of a previously destroyed natural range plant community. This was conservation and wise use management of natural resources. Adams County, Washington. June.

A Range Management professional's observation: Restoration Ecology and Conservation Biology are two disciplines in the biological sciences devoted to conservation of natural resources, especially or mostly of organisms. These subject matter areas, and professionals coming out of them, customarily use some words that differ from those that have been employed by foresters, rangemen, and wildlifers for decades. Most are just old wine in new skins. Even restoration is an old term that refers specifically to that state or level of conservation which is nonuse at the present time and/or for some time in the future for the explicit purpose of building up, bringing back, etc. (ie. restoring) a natural resource that had been at one of those states of resource use known as exploitation or depletion (= exhaustion). All these terms refer to rates of resource use. These latter two rates of use of natural resources are rates that exceed conservation. Rreclamation and rehabilitation are two related terms for principles and practices applied to management of drastically altered, severely disturbed land and associated bodies of water to bring about some improvement from their depleted state.

Restoration, conservation, reclamation, and wise use management were not coined or invented by restoration ecologists and conservation biologists. These professionals took them up from professionals in Agriculture, especially rangemen, foresters, wildlifers, and fisheries scientists who, bolstered with proven practices from agronomists, farmers, and stockmen, have plied the science and art of conservation for years. Welcome aboard newcomers. We old timers in conservation, we the original conservationists, need all the help we can get. Any new perspectives, discoveries, technological advances, etc. that you can offer will be most appreciated. There is more than enough pressing work to go around to conserve and wisely use natural resources, including threatened and endangered species.

Just one thing: we were here first. Please give precedence and priority of use and discovery its proper due. This was our bailiwick first. Restoration ecologists and conservation biologists talk of these things; rangemen, foresters, wildlifers, agronomists, watershed specialists (though they may not have been called that then), even horticulturalists have been doing them for a century or more. Successful grassland, wetland, forest, range, etc. restoration have been hallmarks of Agriculture since conservation came into its own with the Progressive Movement. Restoration of Palouse Prairie through the Conservation Reserve Program and Grazingland Conservation Initiative are two of the more recent examples of this.

Along these same lines, however, it must be emphasized that botanists, zoologists, ecologists, and even natural historians provided much of the basic and applied knowledge that made conservation possible. Even in the first decade or so after World War II jounrals in fields like Ecology, Botany, Entomology, and Natural History provided more practical knowledge in or, at least, relevant to Range Management than much of the contempory stuff in the Journal of Range Management and, now, Journal of Rangeland Ecology and Management.

Recently, some traditional workers in conservation and natural resource management (including supposed rangemen and foresters and even agronomists) have taken to spouting the newer but less applied terms given compliments of restoration ecologists and conservation biologists. Some university units that once proudly proclaimed themselves Departments of Range Management (or Science), Animal and Range Sciences, Forestry and Range Management, or Schools of Forestry now bear such balderdash as "Ecosystem Sciences"'. Fair enough. That is your academic freedom, but only to a point. Academic freedom and institutional effectiveness are not license to come up with Madison Avenue "cutesy" catchphrases in order to "up enrollment"or even try to save dying programs. Professionals put the profession first, not students, clients, patients. Then they let the chips fall they as they will. Range Management is Range Management, Forestry is Forestry, and Agronomy is Agronomy. These discipline-based professions are not "Environmental Science", and trying to pass them off as such is dishonest and a discredit to those agricultural sciences and the professions built upon them.

Reclamation of severely disturbed lands, restoration of depleted ranges, reforestation of cutover forests, and replinishment of game herds (= conservation of natural resources) are in turn the wise application of Agronomy, Range Management, Forestry, Wildlife Management, and so on. These actions and their outcomes are not "Environment Science", Restoration Ecology, or Conservation Biology. Those willing to sell out the traditional agricultural sciences that have enabled resource managers to restore grasslands like those of the Palouse Prairie have a planning horizon that extends only to their retirement (or perhaps only to the next promotion). Time will tell how far "selling your saddle" got you.

As for the rangeman who did the work of this publication it was with professional pride and respect for his profession of Range Management that much of the new jargon was herein pointedly excluded, even rediculed, for what they are: politically expedient, "trendy", "buzz words".

Introduced Species Used as Range In the Palouse Prairie

Many species of grasses and legumes along with some chenopods, crucifers, and composites have been introduced into North America from other continents to be used as"substitutes" for native range plants that for any number of reasons were difficult or even impossible to establish (re-establish) on their original habitats. These alien, exotic, or non-native plant species have been designated by tradition or definition as introduced species or introduced plants (sometimes accompanied with adjectives like forage, feed, wildlife, cover, conservation, etc.). A number of these introduced species were used as both pasture (= tame pasture, introduced pasture, agronomic species used for grazing ) and range (natural pasture, natural grazing land). Most of the introduced plants that were acceptable or successful for range eventually became naturalized, at least to large degree. Many or most of these introduced plant species require human managemaent for initial establishment, but they typically persist indefinitedly once established. Many introduced species have naturalized to the point that once introduced into an area or region they naturally establish themselves.

A number of these introduced plant species were treated separately in this publication with the heading, Introduced, under the biome, Grasslands. A few of the most important introduced species in the Palouse Prairie were also included at the end of this chapter.

166. Crested wheatgrass seeding- Traditionally range has been defined so as to include grazing lands that were reseeded to non-native (= introduced) plants that are managed as if they are native (or naturalized) species and manipulated more by extensive ecological inputs, especially grazing management, than by intensive agronomic methods. The case exemplar for this inclusion of introduced species into the category of range plants and native grazinggrounds was probably the reseeeding of abandoned crop land in the Northern Great Plants and greater Intermountain Region (including the Palouse Prairie) of North America to the Eurasian bunchgrasses known as crested wheatgrass (Agropyron desertorum and A. cristatum).Crested wheatgrass was introduced from Russia and Siberia and used to reseed "go-back land" (abandoned farm land undergoing secondary succession) and marginal crop fields during the first third of the Twentieth Century. These are still, acreage- wise, the major introduced range plants in North America. This is the Berger Tract of crested wheatgrass south of Twin Falls Idaho. October.No FRES or Kuchler designation because this is not native or potential natural vegetation. (Refer to Literature Review for the limitations of limiting the designation of range strictly to indigenous species.) SRM 614 (Crested Wheatgrass).

It should be remembered that some of native range plants do return by secondary succession to lands reseeded to crested wheatgrass. This is especially the case for shrubs like big sagebrush (shown here invading what was once likely a bluebunch wheatgrass-big sagebrush shrub steppe). Rabbitbrush, greasewood, and horsebrush are other examples of the inevitable invasion of woody plants into grasslands in absence of recurrent fire. These shrubs make important contributions to animal diets and they should be recognized as part of the potential natural vegetation (that is, of course, at population levels below those on ecologically deteriorated ranges caused by past cultivation, overgrazing, underburning, etc.).

The contribution of crested wheatgrass to diets of native ruminants like pronghorn antelope (Antilocarpa americana) and mule deer (Odocoileus hemionus) is unfortunately all too often unappreciated by those who decry crested wheatgrass seedings as monocultures of introduced species.

167. Crested wheatgrass range- Crested wheatgrass seen here is just entering dormancy having set seed and completed it's annual growth cycle. This seeding is in southwestern South Dakota (Pennington County). When it is compared with the one in Idaho and the crested wheatgrass specimens in Alberta the reader can grasp the longitidunal range in North America over which this introduced Eurasian species is adapted. July. SRM 614 (Crested Wheatgrass).

168. Crested wheatgrass (Agropyron cristatum)- Crested wheatgrass includes only A. cristatum in the broadest sense. In a more strict or specific sense crested wheatgrass is the generic name for A. cristatum, A,. desertorum and, less commonly, A. fragile.

The definitive authority on crested wheatgrass (in all aspects) is the symposium proceedings edited by Johnson (1986).

169. Stand of immature crested wheatgrass- Crested wheatgrass at the phonological stage known as the boot stage. This is the stage of growth in grasses and grasslike plants when the inflorescence is enclosed in the sheath of the uppermost leaf (all editions of Forages as for eg. Barnes et al., 1995, p. 488). This is often the phenological stage at which herbage or forage (biomass) yield and nutritive value or content (ie. nutrient yield) are at optimum levels, and with respect to each other. Crested wheatgrass is clearly not up to the standard of wheat pasture, but it is good quality by most range (and tame pasture) standards. This example demonstrates why this extraordinary introduced range grass has been so widely planted and how it can complement native range and increase ranch productivity and profitability. Flying D Ranch, Gallatin County, Montana. June. SRM 614 (Crested Wheatgrass).

170. Crested wheatgrass pasture at peak standing crop- This photograph was taken on the same day as the preceding slide, but here on this shallower, more upland land (a bench) crested wheatgrass has advanced to maturity at an earlier date and in this drier than typical spring. (Note the dried, mature seed stalks.) On this less fertile and mesic site this introduced Eurasian grass will continue to survive (including, of course, to reproduce) as naturalized range and serve as what local ranchers call "dryland pasture". (Compare to irrigated pasture on flood meadow three slides below.)

As mentioned for the previous slide, crested wheatgrass does not produce the highest quality grass forage imaginable. Yet, as evidenced by these registered Line One Herefords, diets from crested wheatgrass range are adequate for phenotypic expression of the genetic potential of high performing grazing animals.

No FRES or Kuchler designation. SRM 614 (Crested Wheatgrass). Cooper Hereford Ranch, Gallatin County, Montana. June.

171. Appearance of a crested wheatgrass seeding at end of growing-grazing season- This Clayey range site had been put to the plow and was go-back land when it was reseeded to crested wheatgrass. The seeding has persisted for many years and now also supports scattered native grasses including western wheatgrass, blue grama, and green needlegrass . Perhaps this is one of the lasting "legacies" of this marvelous introduced range grass: it prepared the way for at least partial recovery of the native species. This is the process of reaction (later termed facilitation) in the Clementsian model of plant succession in which each seral stage improves the sere (the total environment of a given site complete with the complete sequence of plant communities that will come to occupy it) for the next seral stage (the plant community at that stage of vegetation development) until the termination at climax. If crested wheatgrass persist indefinitely, as seems likely, as a disclimax or anthropogenic climax it will continue to conserve soil and contribute to range restoration while providing valuable forage for livestock and wildlife. That is a most fitting accomplishment for any pasture crop. The introduction of and development of management programs for crested wheatgrass was an example of Man, Manipulator of Ecosystems, at his best. The establishment of crested wheatgrass range was a technological revolution in grassland agriculture. Pennington County, South Dakota. July. SRM 614 (Crested Wheatgrass).

172. A seeding of crested wheatgrass and a cultivar of dryland alfalfa- This former cropland was seeded to a mixture of crested wheatgrass and a cultivar of the wild, yellow- or variegated-flowered, branch-rooted alfalfa (Medicgo falcata cv. Ladak). The classification by Vallentine (1990, ps. 7-13) of grazing lands as: 1) long-term, 2) medium-term, and 3) short-term was presented in the Introduction of this publication. Crested wheatgrass is the case exemplar of an introduced species being used as or becoming naturalized (vs. native) range. The Vallentine (1990, p. 10) designation of "seeded range (introduced species)" as a category of "long-term grazing land" with duration of stand in excess of 40 years fits crested wheatgrass "to a T". By comparison, persistence of alfalfa is at best of such a short interval that this seeded introduced species would fall into the Vallentine (1990, p. 10) designation of "permanent pasture" under "medium-term grazing lands" (a period of about 10 years). Those working closely with this grass-legume mixture specified however that the Ladak alfalfa is both less palatable and later-maturing (phenological development is slower) than the crested wheatgrass with the result that the latter is grazed heavier (greater degree of use) and earlier in the growing season (Western Section, American Society of Animal Science annual meeting tour, June, 2001). This favors the alfalfa and enables it to remain competitive with the grass. The net result is that Ladak alfalfa is not grazed out and the seeding is not converted to a single species stand of crested wheatgrass. Alfalfa has also been grown in seedings with intermediate wheatgrass (Agropyron intermedium), Russian wild rye (Elymus junceus= Psathyrostachys juncea ), and in mixtures of these two species plus crested wheatgrass (Heinrichs, in Campbell and Herbel, 1975, ps. 54-56). No FRES or Kuchler designation. Variant of SRM 614 (Crested Wheatgrass). Gallatin County, Montana. June.

173. Dryland creeping alfalfa – An individual plant of Ladak alfalfa (Medicago falcata cv. Ladak). This is a variety of the wild yellow-flowered alfalfa that is native to Eurasia. Ladak was selected from among native populations of M. falcata in the province of Ladakh, India (Heinrichs, 1963, ps. 317-320 passim). Ladak has variegated inflorescences and a branched-root arrangement (vs. the more pronounced or typical pattern of the dicotyledon taproot system). This cultivar (accession may be the more precise designation) was selected for rainfed agriculture in a semi-arid region and it has been used in intercrosses with other M. falcata lines to produce hybrid types having creeping rootstocks (creeping-rooted alfalfa) (Heinrichs, 1963, p. 322-324).

Alfalfa is one of man's most important agronomic forage legumes, especially as a hay crop. Most cultivars of common alfalfa (M. sativa) can be grown without irrigation only in sub-humid or wetter regions. There are a few alfalfa cultivars that have been selected for dryland agriculture in semi-arid regions. These are generally either M. sativa X M. falcata hybrids or M. falcata selections. Ladak is one of the latter which holds some promise for seeded range. Gallatin, County, Montana. June.>

174. Alfalfa- Leaves and inflorescences of Medicago sativa. Numerous Medicago species are of economic importance as pasture and hay crops. Alfalfa is the most important of all. It is the most important hay species in California, the agricultural wonder of the world. It is probably also the most important hay crop in the Agricultural Region known as the Grazing and Irrigated Crops Region traditionally viewed as extending from the Front Range of the Rockies (western edge of the Great Plains) westward to the crest of the Sierra Nevada and Cascade Ranges. While there are important introduced grass species (eg. timothy, redtop, reed canarygrass) grown for hay in various locations throughout these regions alfalfa is likely the most important hay crop in the vast area known as the Western Range.

Development of alfalfa cultivars which are adapted for either dryland pasture or even range has long been a goal of plant breeders in the Western Range. Unfortunately this remains a largely unfulfilled goal, but some breeders have developed cultivars of the creeping (= creeping-rooted) alfalfa. Whereas the typical M. sativa varieties have a prominent tap-root system the wild yellow-flowered alfalfa (M. falcata) has a "branched-root" form of the fundamental dicot taproot system. This morphological shape or pattern of root arrangement permits this Medicago species to exhibit a "creeping" or "spreading" habit rather than the bunched or tufted habit of typical M. sativa plants. This creeping rootsystem permits expansion by asexual reproduction (and consequent better survival under grazing) in M. falcata. Alfalfa breeders have developed several M. falcata X M. sativa hybrid (sometimes identified as M. media) cultivars including Rhizoma, Rambler, Teton, Nomad, Roamer, and Drylander with the latter being the one best adapted to dryland pasture production. Some of these have creeping rootstocks or even rhizomes under certain conditions (eg. Rhizoma, Nomad, Rambler). Unfortunately these are often low-yielding and relatively restricted in their range of adaptation (see review by Heinrichs, 1963 and Heinrichs, in Campbell and Herbel, 1975, p.54-57). Overall these cultivars are relatively drought-tolerant and persistent on seeded range, at least by legume standards, but the goal of legumes having persistence on range like that of introduced grasses is not yet a reality.

Alfalfa, like the other major agronomic legumes, is a member of the Papilionoideae (= Faboideae) subfamily). Papilionaceous legumes have a corolla consisting of five petals: the largest is a single petal known as the standard or banner, two central petals fused as the keel, and two "side" (lateral) petals known as wings. The major agronomically important nodulated legumes— those entering into mutualistic symbiosis with nitrogen-fixing bacteria like Rhizobia species which form nodules on the host roots— are all in this subfamily, the Papilionoideae.

175. Desert crested wheatgrass (Agropyron desertorum) range- Desert crested wheatgrass instead of "regular" crested wheatgrass is grown in drier areas to which this group of Eurasian perennial bunchgrasses is adapted. This desert crested wheatgrass stand was planted in the High Lava Plains province (Franklin and Dyrness, 1973, p.6) of southcentral Oregon. The native Wyoming big sagebrush had re-established from the soil seedbank in this seeded single-species stand of naturalized bunchgrass to provide an outstanding example of the "right blend" of crested wheatgrass and big sagebrush. Physiognomy, structure, and species composition of this naturalized range was "danged near perfect".

Harney County, Oregon. June. Estival aspect; peak standing crop. No FRES number or Kuchler unit for this non-native (but naturalized) range cover type. SRM 614 (Crested Wheatgrass).

176. Naturalized range of desert crested wheatgrass- These two views of the same range (a different range from that presented in the preceding photogarph) that developed from a seeding of desert crested wheatgrass provided the student with an all-inclusive picture of "the way a crested wheatgrass range should look". The single-species stand of this Eurasian bunchgrass had been invaded by scattered plants of Wyoming big sagebrush resulting in the species composition, community structure, and, consequently, proper ecosystem function of one of the single most important and valuable cover types of rangeland in parts of the Northern Great Plains and Intermountain Region of North America.

At various points in this publication beginning students in Range Management have been advised to familarize themselves with the classification of grazing lands based on longevity or duration of pasture developed by Vallentine (2001, ps. 7-13). Vallentine's treatment was comprehensive and covered all grazing lands, both natural or native and introduced or agronomic pasture (a dichotomy that Vallentine [2001, ps.7, related 15-16] showed to be unreliable for certain uses). The Vallentine classification of grazing lands is, in the current author's experience, the most reliable, comprehensive, and useful of any other such treatment of grazing lands (= grazinglands as one word, if the reader prefers).

In the comprehensive, pasture longevity-based system of Vallentine (2001, ps. 7-13) crested wheatgrass grazing land falls under the categories of "long-term grazing land (synonym range), seeded range (introduced species)". Simply put, crested wheatgrass is introduced range. In fact, crested wheatgrass was probably the single most common example of why the term range was first officially defined by the American Society of Range Management (Huss, 1964) to include "... lands that are revegetated naturally or artificially to provide forage cover that is managed like native vegetation".

Technical note on syntax of the language: It is redundant and affected usage to use the words "native range" except in context of drawing the clear distinction between native range vs. introduced range where the latter is the shortened version of the Vallentine (2001, p. 8) term of "seeded range (introduced species)". If one is comparing or distinguishing between range made up of seded introduced species and range comprised of seeded native species or contrasting seeded introduced range to native (non-seeded) range, the use of native range is quite appropariate. Otherwise it is inappropriate and marks one as a novice, pedant or greenhorn. For example, to speak of sagebrush steppe or tallgrass prairie as "native range" without contrast or comparison to, say, crested wheatgrass range or weeping lovegrass (Eragroistic curvula) range is stilted, redundant, and unnecessary verbage. In context of the latter examples, the foolish use of "native range" is the equivalent of describing livestock as "domestic livestock" (without contrast to feral livetock). Use the language of our discipline and profession properly so as to foster respect for them, or do not use our "lingo" at all.

Crested wheatgrass (both A. desertorum and A. cristatum) have proven to be some of the useful non-domesticated plants introduced into North America. These two species have been shown time and again to be invaluable for soil and water conservation and wildlife habitat (feed and cover) as well as in provision of forage for livestock.

Harney County, Oregon. June. Estival aspect

177. Individual plants of desert crested wheatgrass (Agropyron desertorum)- These two robust and ungrazed specimens of desert crested wheatgrass provided examples their species and illustrated cespitose morphology of bunchgrasses in general. Once individual plants have developed to this stage of maturity and have aquired this extreme degree of rank growth they are no longer palatable to grazing animals (unless animals are feeling severe "hunger pangs" if not suffering from "holler belly").

These two grass plants were chosen from ungrazed areas to give viewers "purple ribbon" textbook examples of this species. In a grazed pasture an individual such as either of these "grand champions" is known as a wolf plant: "1) an individual plant that is generally considered palatable, but is not grazed by livestock, 2) an isolated plant growing to extraordinary size, usually from lack of competition or utilization" (Bedell, 1989).

Harney County, Oregon. June. Peak standing crop; soft-seed phenological stage.

178. Spikes of desert crested wheatgrass- Inflorescences of grasses in the Tritaceae or, also, Hordeae (wheat or barley tribe) have inflorescences known as spikes which are distinguished by having spikelets sessile (without a pedicel or short "stalk") on the rachis (central axis of a grass inflorescence).

Harney County, Oregon. June. Soft-seed phenological stage.

179. Intermediate wheatgrass in the Palouse Prairie Region- A former native bunchgrass steppe in heart of the Palouse Hills was converted into this dry-land pasture of intermediate wheatgrass with substantially greater grazing capacity. Another melanchology instance of what Charlie Russell dubbed "Trails Plowed Under" as the native grassland gave way to more agriculturally productive --and profitable-- tame (ie. agronomic) pasture.At least it was still grazing land. Furthermore, in this instance the introduced forage grass provided complementary pasture for adjoining cropland also used for grazing (eg. small grains pasture, crop aftermath) or harvested forage (primarily hay) along with some scattered remnants of Palouse Prairie range.

This was also an example of wise and efficient use of water. This tame pasture was being managed strictly as a rain-fed forage crop even though this field was bottomland with fertile soil such that domesticated, high-yielding crops would respond faovrably (physiologically if not profitably) to irrigation. In such a grassland agriculture management system limited water and expensive fossil fuels could be used to greater advantage on other fields for other crop species. Utilization of fields like this for higher-producing agronomic forages also enabled puirebred beef cattle to express their genetic potential which could then be evaluated. Such evaluation for genetic potential (eg. progeny testing for economically important traits) would not be possible (at least not to such extent) on natural pastures like the very limited remaining Palouse Prairie range that in such a fertile area was limited to steeper or rougher land which was marginal for field crops like small crops and oil seeds. An example of wise use management of financial and cultural resources as well as natural resources.

Whitman County, Washington. June, early hard dough stage of grain.

180. Robust individual of intermediate wheatgrass- The cespitose morphology of intermediate wheatgrass was shown to good advantage in this "hale and hearty" specimen that was growing on the dry-land pasture presented in the immediately preceding photograph. Whitman County, Washington. June, early hard dough stage of grain.

181. Spikelets up close- Spikelets of intermediate growing on the non-irrigated, bottomland pasture showed and described above. Whitman County, Washington. June, early hard dough stage of grain.

182. A plant breeding- Spikelets at anthesis on spike of intermediate wheatgrass. Big Horn County, Wyoming. Late June (earlhy summer); anthesis.

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