True Prairie - IB

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

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

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

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

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

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

Theodore Roosevelt National Park (North Unit), McKinzie County, North Dakota. Late June, early estival aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-59 (Wheatgrass-Needlegrass). Savanna variant of SRM 607 (Wheatgrass-Needlegrass) and SRM 610 (Wheatgrass). Cold Temperate Grassland 142, Plains Grassland 142.1 but no appropriate series in Brown et al. (1998). Northwestern Great Plains- Missouri Plateau Ecoregion 43 a (Bryce et al., Undated).

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

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

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

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

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

Theodore Roosevelt National Park (North Unit), McKinzie County, North Dakota. Late June, early estival aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-59 (Wheatgrass-Needlegrass). Savanna variant of SRM 610 (Wheatgrass). Cold Temperate Grassland 142, Plains Grassland 142.1 but no appropriate series in Brown et al. (1998). Northwestern Great Plains- Missouri Plateau Ecoregion 43 a (Bryce et al., Undated).

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

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

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

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

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

Theodore Roosevelt National Park (North Unit), McKinzie County, North Dakota. Late June, early estival aspect. Green needlegrass was in milk- to soft-dough grain stage. FRES No. 38 (Plains Grasslands Ecosystem). K-59 (Wheatgrass-Needlegrass). Savanna variant of SRM 607 (Wheatgrass-Needlegrass). Cold Temperate Grassland 142, Plains Grassland 142.1 but no appropriate series in Brown et al. (1998). Northwestern Great Plains- Missouri Plateau Ecoregion 43 a (Bryce et al., Undated).

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

126. Long clusters of a shortgrass- Panicles of Sandberg's bluegrass that was growing on a true prairie range in the sedimentry Northern Great Plains. Garfield County, Montana. Mid-June; soft-dough grain stage.

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

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

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

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

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

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

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

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

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

The following short section presented some papilionaceous legume that are forbs on various rangeland cover types of both true prairie prairie and mixed prairie in the Northern Great Plains. Specimens shown here were growing on various ranges (pastures) varying in range condition from Excellent down to Good through Fair ending with Poor condition class. These leguminous forbs were all in tribe, Galegeae except for one species that was in Hedsareae tribe.

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

Standard reference: the old standby for treatment of the Astragalus species on the Western Range was Hermann (1966, ps. 35-79) who organized species according to subgenera or sections. Also recommended, especially for color plates, was Rickett (1973, ps. 410-415 for plains species). For taxonomic treatment of Astragalus species in the general Great Plains (except for Southern High Plains) Region was Great Plains Flora Association (1986, ps. 421-436)

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

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

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

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

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

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

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

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

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

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

Garfield County, Montana. Late June.

Standard reference: the old standby for treatment of the Oxytropis species on the Western Range was Hermann (1966, ps. 118-127). For plains species (minus some of the Southern Great Plains in Texas) see Great Plains Flora Assocition (1986, ps. 467-471).

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

Garfield County, Montana. Mid-June.

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

and

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

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

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

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

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

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

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

154. Cryptogams on wheatgrass-needlegrass range- Local colony of lesser spikemoss (Selaginella densa) with some plants of tumbleweed shield lichen (Parmelia chlorochroa= Xanthoparmelia chlorochroa) lichen on a western wheatgrass, thickspike wheatgrass, needle-and-thread-Sandberg's bluegrass prairie range on the Northern Great Plains. This species of lichen was implicated in elk poisoning in Wyoming. P. chlorochroa has been recognized as an important cryptogam throughout much of the Northern Great Plains (Maccracken et al., 1983). This natural pasture was in Good range condition class. Some culms of Sandberg's bluegrass were present.

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

Garfield County, Montana. Mid-June.

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

Garfield County, Montana. Mid-June.

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

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

Garfield County, Montana. Mid-June.

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

Garfield County, Montana. Mid-June.

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

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

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

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

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

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

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

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

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

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

Transition Grassland of the True Prairie

There was no more difficult (and, perhaps, dubious) distinction between the grassland associaions of Clements (1920) than that of mixed prairie versus true prairie. This was not always a tenuous distinction, but it was downright arbitrary in certain areas where adjoining soil series or soil associations resulted in dramatically different--at least to the experienced "range eye"--grassland plant communities.

Shown below was one such example of a climax grassland community co-dominated by needle-and-thread and western wheatgrass. That combination was of itself unique because needle-and-thread thrives on sandy soils like those of some sandhills range sites whereas western wheatgrass does best on finner-textured (clay) soils. Soil texture classes like sandy clay or sandy clay loam provided ideal edaphic habitats for this combination of two cool-season grass species.

Given the arbitrary distinction of mixed prairie versus true prairie, this example was included in two Grassland chapters entitled 1) True Prairie and 2) Mixed Prairie-IA, Southern and Central Great Plains.

167. Two midgrasses and one herd of stockers- True or mixed prairie (depending on interpretation or perception) in the Central Great Plains co-dominated by western wheatgrass and needle-and-thread with blue grama as the associate. Also a "tad" of buffalograss. Even fewer plants of threadleaf caric sedge and sixweeks fescue. Main forb was wavyleaf thistle (first and third of these slides). Widely scattered plants of sand sagebrush.

The climax range vegetation was in the vernal society part of the annual cycle. It was at peak standing crop (or just past) of the two dominant cool-season, festucoid grasses. Some shoots of both co-dominants had turned the amber of their completed growth cycle for this year while other shoots were in progress to that growth or phenological stage. In the estival society phase to follow blue grama would mature to its peak standing crop, but this would be less than than that of the cool-season co-dominant grasses.

These three slides presented the landscape (and landscape-scale) along with physiogonomy of this climax range plant community (Excellent range condition class) which was being grazed by stockers (all steers).

Washington County, Colorado. Mid-June; late vernal aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-59 (Wheatgrass-Needlegrass). SRM 607 (Wheatgrass-Needlegrass). Loamy plains range site (Soil Conservation Service, 1986). High Plains- Flat to Rolling Plains Ecoregion, 25d (Chapman et al., 2006).

168. Range vegetation and range cattle up close- Three views of the internal structure and species composition of a climax true or mixed prairie (depending on interpretation of the viewer) in the Central Great (High) Plains. Needle-and-thread and western wheatgrass were the co-dominants while blue grama was the associate species of this Excellent range condition class grassland. Samd sagebrush (center foreground of third slide) added a woody (shrub) component but it was nowheree abundant to comprise a woody layer. Likewise the few widely scattered plants of plains pricklypear did not a shrub layer make. The principal (the only) forb was wavyleaf thistle (see the third of three slides in the immediately preceding slide-caption set).

The crossbred steers (roughly 650 pounds) were extremely gentle and curious. Steers came up to within two feet of the photographer, but would not lick his extended hand (these slides were taken using a 28mm lens so cattle were considerably closer than they appear). This author has found--in the vast majority of instances--that docile cattle and high-quality feed, high-successional stage range go hand-in-glove. The obvious explanation is that good husbandry, sound stewardship includes all range resources. This includes human and financial resources, but the author was not privileged to partake of these latter two categories.

Nice story. "All lived happily after ever" (at least until harvest time and end of the life cycle).

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 the Palouse 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 Palouse Prairie, given that it is primarily a representative part of the bunchgrass prairie or Agropyron-Stipa Association (Clements, 1920, 149-152). With absence of Bouteloua species and buffalograss (Buchloe dactyloides) this author could not in his rangeman's mind justify this bunchgrass prairie (steppe) as a form of mixed prairie, the Stipa -Bouteloua Association (Clements, 1920, ps. 135-139) though it obviously shares an affinity with it and the true prairie. It had a much greater vegetational relatedness to true prairie than mixed prairie based on physiogonomy and cover composition by the same dominant climax grass species.

169. 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) form 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 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).

170. 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).

171. 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.

172. 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.

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

Crook County, Wyoming. Mid-June.

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

Crook County, Wyoming. Mid-June.

179. Now they're ready- Mature panicles of green needlegrass just before grain shatter. Musselshell County, Montana. Mid-June (late spring); grain-ripe stage sust before seed-shatter.

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

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

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

Powder River County, Montana. Mid-June; late vernal aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-57 (Grama-Needlegrass-Wheatgrass). SRM 606 (Wheatgrass-Bluestem-Needlegrass). Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40). Northwestern Great Plains-Montana Central Grasslands Ecoregion 43n (Woods et al., 2002).

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

Low, sawthooth hills behind were a Breaks range site.

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

Northern True Prairie Savanna or Northern Mixed Grass-Shrub Steppe

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

207. Mixed prairie savanna- Savanna form of mixed prairie co-dominated by needle-and-thread and prairie sandreed with locally heavy cover from large plants of silver sagebrush. This savanna developed on the immediate floodplain (first terrace) of the Little Missouri River in the Badlands formed by this river. The associate herbaceous species on this river-bottom range was needle spikerush (Eleocharis acicularis) which was shown in foreground in the second slide. Another local associate species was smooth scouring-rush or smooth horsetail (Equisetum laevigatum).

Trees in background were a floodplain forest of eastern cottonwood (Populus deltoides subsp.monilifera), peachleaf willow (Salix amygdaloides), and green ash (Fraxinus pennsylvanica). This forest cover type was covered in the chapter Miscellaneous Forest under Woodlands and Forests.

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

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