Mixed Prairie - ID

Northern Great Plains

Mixed prairie comprises the vast majority acreage of Great Plains grasslands. Mixed prairie was named for its most obvious floristic aspect: it is comprised of tall-, mid-, and shortgrass species. This mixed composition is not always present as there are posclimax tallgrass-dominated communities, even consociations of tallgrass species (eg. sand bluestem [Andropogon hallii]), on deeper and or sandy soils, preclimax shortgrass (eg. buffalograss [Buchloe dactyloides]) communities on shallow soils and generally more xeric range sites, and yet other grassland communities (usually consociations) made up almost exclusively of a single species (eg. western wheatgrss [Agropyron smithii] on swale range sites). Sometimes speakers will refer to mixed prairie as "midgrass prairie" to be consistent with tallgrass prairie and shortgrass plains and reflecting an overall physiogonomy that is intermediate (= "midway") between the latter two major forms of central-continent grasslands.

The ultimate reference for mixed prairie (and shortgrass prairie) from standpoint of descriptions of range vegetation and response to disturbance is--and will likely always be--the classic Grasslands of the Great Plains by Weaver and Albertson (1956). Definitive source of knowledge regarding mixed paririe from an ecosystem perspective is that of Coupland (in Coupland, 1992, ps. 151-182). The comprehensive monograph on shortgrass plains (=prairie, steppe) edited by Lauenroth and Burke (2008) overlapped--by necessity--parts of the mixed prairie so as to be a valuable reference for western extremities of mixed prairie.

Northern Great Plains Mixed Prairie

The huge complex of grasslands in the greater Northern Great Plains Region (including margins of adjacent physiographic provinces) consist of a mosaic of cover (dominance) types. The definitive source for this region was Barker and Whitman (1989), the vegetational units of which were incorporated into Society for Range Management rangeland cover types (Shiflet, 1994). In the section below various of these range cover types were presented and described. Other rangeland cover types (Shiflet, 1994) in the Northern Great Plains (Barker and Whitman, 1989) fit the textbook designation of true prairie and were included in the chapter of that title under the Grasslands heading. Also included in the chapter, True Prairie, was a vegetational mosaic of ponderosa pine (Pinus ponderosa) forest and true prairie grasslands.

General comment on organization: Arrangement of vegetational units in chapters and sub-chapters of Range Types of North America was based on classification and description of vegetation as established by such seminal works as Clements (1920), Weaver and Clements (1929, 1938), and Braun (1950) and not on (or according to) geographic region or physiographic province as was followed (sort of ) for forest cover types (Eyre, 1980) and rangeland cover types (Shiflet, 1994). In Range Types range vegetation was organized as to climax or potential units of natural plant communities at organizational levels extending from biomes or formations, associations, etc. down to range types and, in some cases, range sites. Certain of the major and more diverse associations (mostly Clementsian associations) or major subunits of biomes (such as mixed prairie in this instance) were subdivided into chapters organized by physiographic provinces or related natural geologic units (eg. Northern, Central, Southern Great Plains). Again, however, it was vegetation--not region or geologic province--that served as primary or fundamental basis of distinction and arrangement of chapters within this publication.

Further note on organization and location: In addition to the preceding note as to location of true prairie range types, the reader was directed to the chapter, Southern and Central Forest, under the Woodlands and Forests heading for treatment of Rocky Mountain juniper (Juniperus scropulorum) woodland and green ash (Fraxinus pennsylvanica) open forest types of the Northern Great Plains. These forest cover types had developed within the surrounding regional or zonal grassland climax covered immediately below and in the True Prairie chapter.

1. A textbook example- Mixed prairie on the unglaciated (sedimentary) Northern Great Plains. The potential natural (climax) vegetation at zonal scale across this part of the Great Plains was the wheatgrass-needlegrass cover type, but due to an absorptive clay, a clayey soil over acid clay shale, (Soil Conservation Service, 1976) the range environment of the grassland shown was more mesic and generally less harsh than habitats more typical of this region. In this regard it was most readily regarded as an edaphic climax (the polyclimax theory of Arthur Tansley). The climax range plant community for this range site (Porous Clay) was mixed prairie consisting of species of : 1) tallgrass (prairie sandreed, big bluestem, switchgrass), 2) midgrass (western wheatgrass, tall dropseed, little bluestem), and 3) shortgrass (blue grama). Throughout much of this region, especially farther north and west, little bluestem grows only to a shorter mature height than farther to the south and east so that it frequently is more of a mid- than the typical tallgrass species. In the range plant community described here little bluestem could be regarded as either a tallgrass or, alternatively, as a midgrass.

Western wheatgrass was the dominant cool-season grass in this climax range vegetation. There was some cover of needle-and-thread and green needlegrass, two other cool-season native grasses in this climax range vegetation. Cover, as well as density, these two Stipa species amounted only to Trace amounts. Naturalized Kentucky bluegrass was present at an even smaller proportion of total plant cover. This range plant community was clearly comprised primarily of warm-season grass species. This grassland was, however, mixed prairie from the perspective of having both warm- and cool-season species as well as in the strict meaning of having dominant grasses of more than one life-or growth-form height.

There were a few plants of sun caric sedge (Carex heliophila= C. inops subsp. heliophila= Carex pensylvanica). Forbs were not common on this range, but there were scattered plants of prairie-turnip, breadroot scurf-pea, or Indian breadroot (Psoralea esculenta), western yarrow (Achillea lanulosa= A. millefolium subsp. lanulosa), pale agoseris or pale false dandelion (Agoserus glauca), plains or prairie spiderwort (Tradescantia occidentalis). There were also plants of prairie rose (Rosa arkansana) which could be viewed as either forb or shrub. Though not common there were numerous plants of the shrub, silver or dwarf sagebrush (Artemisia cana).

Dominant and associate status of plant species varied considerably over remarkably short distances due to the patchwork of local habitats (microsites) on this land. The small-scale size of these microhabitats was undoubtedly due to numerous variables, but one of the most obvious was soil depth (or, more obvious, depth of depressions in land surface) which was attributable in some large degree to erosion (much of this soil loss appeared to have been geologic rather than accelerated erosion). This was shown to better advantage in the second of these two slides and in the first slide in the next succeeding set of two slides and their caption.

The soil of this Porous Clay range site was the series, Graner clay (Fine, Montmorillonitic, Acid, Mesic Ustic Torriorthents) that can extend to depths of 60 inches (Soil Conservation Service, 1976; 1990). This soil is highly susceptible to erosion, by both water and wind (Soil Conservation Service, 1976). Several shallow, though wide, gullies were present on this high Good to Excellent condition class range. Most of the herbage in these gullies was that of western wheatgrass. This festucoid grass was the dominant cool-season species of this range polant community and the grass species most advanced in this estival plant society. Hence the prominent green color in lower-lying depressions and gullies. Western grass was apparently better adapted to either shallower soils of gullies or, alternatively, to microsites that ponded or retained water--perhaps more water--longer. By contrast, the tallgrass species (primarily prairie cordgrass, prairie sandreed, and switchgrass) grew on higher ground (small parcels of land having slightly higher elevations of soil surfaces and, presumedly, deeper soil profiles).

Butte County, South Dakota. Mid-June; late vernal aspect (warm-season season grasses had only early growth). FRES No. 38 (Plains Grasslands Ecosystem). K-59 (Wheatgrass-Needlegrass). SRM 608 (Wheatgrass-Grama-Needlegrass). Cold Temperate Grassland142- Plains Grassland 142.1, without an Series designation in Brown et al. (1998, p. 40). Porous Clay (Graner clay) range site (Soil Conservation Service, ). Northwestern Great Plains- Semiarid Pierre Shale Plains Ecoregion 43g (Bryce et al., Undated).

2. In one of its purest forms- Northern Mixed Prairie of tallgrass, midgrass, and shortgrass species along with a few forbs and a shrub component of silver sagebrush.. This mixed prairie developed on a clay shale in the unglaciated (sedimentary) part of the Northern Great Plains. This grassland community was best viewed (explained) from the Tansian polyclimax theory as an edaphic climax. Major grasses in this range vegetation included prairie cordgrass, prairie sandreed, little bluestem, switchgrass, western wheatgrass, tall dropseed, and blue grama. Dominance and associate status of these species varied greatly over short distances due to the microhabitats on the land surface due in large part to differential erosion of soil (below in this caption). This seemed to have been geologic rather than accelerated erosion.

Western wheatgrass was the dominant cool-season grass in this climax range vegetation. There was some cover of needle-and-thread and green needlegrass, two other cool-season native grasses in this climax range vegetation. Cover, as well as density, these two Stipa species amounted only to Trace amounts. Naturalized Kentucky bluegrass was present at an even smaller proportion of total plant cover. This range plant community was clearly comprised primarily of warm-season grass species. This grassland was, however, mixed prairie from the perspective of having both warm- and cool-seaon species as well as in the strict meaning of having dominant grasses of more than one life-or growth-form height. In the range plant community of this mixed prairie there were climax species of 1) tallgrasses (prairie sandreed, prairie cordgrass, switchgrass); 2) midgrasses (little bluestem, western wheatgrass, tall dropseed), and 3) shortgrasses (blue grama was the major one)..

There was some cover of sun caric sedge, but this ws limited. Forbs included Indian breadroot or breadroot scurf-pea, western yarrow, and plains or prairie spiderwort. Silver sagebrush was the main shrub although prairie rose could be regarded as shrub or forb.

This seemed certainly to be an ecotonal or transitional grassland between the tallgrass prairie of the Nebraska Sandhills and true prairie farther to the north and west as well as less mesic range sites in this general immdediate area. It was a transition grssland between more mesic environments of the subhumid zone to the east and the less mesic habitats even farther west in the semiarid precipitation zone.

Dominance of plant species varied primarily based on either soil depth or depth of depressions in the land surface. The soil of this Porous Clay range site was the series, Graner clay (Fine, Montmorillonitic, Acid, Mesic Ustic Torriorthents) that can extend to depths of 60 inches (Soil Conservation Service, 1976; 1990). This soil is highly susceptible to erosion, by both water and wind (Soil Conservation Service, 1976). Several shallow, though wide, gullies were present on this high Good to Excellent condition class range. Most of the herbage in these gullies was that of western wheatgrass. This festucoid grass was the dominant cool-season species of this range polant community and the grass species most advanced in this estival plant society. Hence the prominent green color in lower-lying depressions and gullies. Western grass was apparently better adapted to either shallower soils of gullies or, alternatively, to microsites that ponded or retained water--perhaps more water--longer. By contrast, the tallgrass species (primarily prairie cordgrass, prairie sandreed, and switchgrass) grew on higher ground (small parcels of land having slightly higher elevations of soil surfaces and, presumedly, deeper soil profiles).

This phenomenon was shown to better advantage in the first of these two slides and in the second slide in the immediately preceding set of two slides and their caption. As described above there was some cover (Trace amounts) of two other cool-season native grasses ,needle-and-thread and green needlegrass, in this climax plant community as well as even less cover of the ecological invader, Kentucky bluegrass.

3. All the components were there - Two "photoquadrants" provided a "who's who" of range plant species on Northern Mixed Prairie that developed on soil derived from Pierre Shale parent material. Plant species identifiable within frames of these photographs included: prairie cordgrass, prairie sandreed, switchgrass (tallgrasses); little bluestem, tall dropseed, western wheatgrass (midgrasses); blue grama (shortgrass), sun caric sedge (grasslike plant); plains spiderwort, pale false-danedline, breadroot scurf-pea or prairie turnip, western yarrow (forbs); silver sagebrush and prairie rose (shrubs or shrublike plants). Several shoots of silver sagebrush were standing prominently in center foreground of the first slide. The taller dead shoots (straw of last year's shoots) were mostly of prairie cordgrass and prairie sandreed with fewer shoots switchgrass.

Warm-season grasses, especially tallgrass species, were in early stages of annual shoot growth and appeared most conspicuously as tan-, brown-, or amber-colored leaves and culms of last summer's herbage. Most of the green shoots in these photographs were those of western wheatgrass, the dominant cool-season species on this range. There was some cover the cool-season Stipa species, needle-and-thread and green needlegrass, but only at levels best described as Trace. There was even less cover of the naturalized cool-season Kentucky bluegrass. This climax range vegetation was made up overwhelmingly of warm-season species, and it was a "prime" textbook example of mixed prairie in the unglaciated Northern Great Plains, a mixed prairie community consisting of: 1) tallgrasss, 2) midgrass, and 3) shortgrass species.

This phenomenon was shown to better advantage in the second and in the first slide of the two slides- caption that preceded this set of two slide-caption unit. There was some cover of needle-and-thread and green needlegrass, two other cool-season native grasses in this climax range vegetation. Cover, as well as density, these two Stipa species amounted only to Trace amounts. Naturalized Kentucky bluegrass was present at an even smaller proportion of total plant cover. This grassland was, however, mixed prairie from the perspective of having both warm- and cool-season species as well as in the strict meaning of having dominant grasses of more than one life-or growth-form height.

4. A synopsis view- Edge of a relatively wide yet shallow gully and adjoining land with slightly higher soil surface (due to less soil loss via erosion) on Northern Mixed Prairie in the unglaciated (sedimentary) Northern Great Plains. Parent material of this soil was Pierre Shale. The soil of this Porous Clay range site was the series, Graner clay (Fine, Montmorillonitic, Acid, Mesic Ustic Torriorthents) that can extend to depths of 60 inches (Soil Conservation Service, 1976; 1990). This soil is highly susceptible to erosion, by both water and wind (Soil Conservation Service, 1976). Several shallow, though wide, gullies were present on this high Good to Excellent condition class range.

Most of the herbage in these gullies was that of western wheatgrass, the dominant cool-season grass in this range plant community and thus the species that furnished most of the green herbage in this estival plant society. Western grass was apparently better adapted to either shallower soils of gullies or, alternatively, to microsites that ponded or retained water--perhaps more water--longer (There was some cover of needle-and-thread and green needlegrass, two other cool-season native grasses in this climax range vegetation. Cover, as well as density, these two Stipa species amounted only to Trace amounts. Naturalized Kentucky bluegrass was present at an even smaller proportion of total plant cover.) This grassland was, however, mixed prairie from the perspective of having both warm- and cool-season species as well as in the strict meaning of having dominant grasses of more than one life-or growth-form height.

Most of the grass with tan- or light-brown colored straw (last year's shoots) was little bluestem, which is best regarded as a midgrass species in this part of the Northern Great Plains. Such patches or local populations of little bluestem were almost single-species stands. These constituted little bluestem consociations. The broader-leafed shoots in immediate foreground were of prairie sandreed produced in the previous (last) summer. Some of the grass cover that was growing intermingled with prairie sandreed in this local microhabitat was switchgrass. Most of the taller green shoots in this vernal society were, as explained in the preceding paragraph, those of western wheatgrass.

Silver sagebrush was the major surub species on this range.

5. A nice mix- Two slightly different views of mixed prairie in Excellent (at least, high Good) range condition class in the unglaciated (sedimentary) part of the Northern Great Plains. The major grass species in these slides were growing on the banks (edges) of gullies. The larger plants with taller shoots were prairie cordgrass and prairie sandreed. There were numerous green shoots of western wheatgrass, the major (and, sometimes, local dominant) cool-season species. There were also dead sexual shoots (complete with grain-shed spikes) of western wheatgrass along with dead (last year's) sexual shoots of tall dropseed in the foreground of both of these slides. Silver sagebrush--barely visible in midground of slides--was the major shrub speices. Most of the broad-leafed plants were prairie rose. There were also some plants of the monocotyledonous and dicotyledonous forbs, plains or prairie spiderwort and Indian breadroot or breadroot scurf-pea, respectively.

This climax range plant community developed on soil derived from Pierre Shale parent material. Soil series was Graner clay (Fine, Montmorillonitic, Acid, Mesic Ustic Torriorthents). Profile of this series can extend to depths of 60 inches (Soil Conservation Service, 1976; 1990). Due to its high content of finely textured clay this soil is highly prone to erosion by both wind and water. This feature was shown in these photographs of the land surface at edge of a shallow gully. These photographs were taken under a heavy overcast sky so there was no glare on the surface of this Graner clay series.

It was shown in preceding slides that most of the herbage in such gullies was of western wheatgrass, the dominant cool-season species of this climax range vegetation. It was explained above that dominance (and status of associate species) varied among the several grass species with the tallgrass species dominating (dominance varying species-by-species over comparatively short distances) on non-gullied (higher ground levels) land which presumedly had deeper soil. Western grass was apparently better adapted to either shallower soils of gullies or, alternatively, to microsites that ponded or retained water--perhaps more water--longer.

6. A little silver- Leaders of a small plant of silver sagebrush that grew on a mixed prairie grassland, Porous Clay range site (Graner clay soil series), in unglaciated Northern Great Plains. This slide presented features of leaves, especially leaf arrangement, on upper shoots of silver sagebrush which is a major shrub over much of the Northern Great Plains. The plant featured here was part of the range plant community described immediately above.

Silver sagebrush has much variability as to plant size and shape ranging from small stunted-appearing (like this cute little guy) to giants of their race. An example of these good ole big 'uns was shown below in the section devoted to grass-shrub savannahs in Northern Great Plains.

Butte County, South Dakota. Mid-June; early stage of annual growth.

7. Across the breaks- Physiography of Cheyenne River breaks in the Northern Great Plains. The first of these two slides was landscape-scale view showing mixed prairie grassland and a local green ash woodland and stand of Rocky Mountain juniper in near background. Second slide was a closer view showing near-physiography, including a hogback (left half of this photograph), and the green ash and Rocky Mountain juniper stands (center near-background). A hogback is a geologic term defined by the American Geological Institute as indicating "a ridge with a sharp summit and steep slopes of nearly equal inclination on both flanks…"; more specifically " a long, narrow sharp-crested ridge formed by the outcropping edges of very steeply inclined or highly tilted resistant rocks … produced by differential erosion" (Gary et al, 1972). An isolated "tooth" or "spine" along the hogback may resemble a small butte such as that in left midground of the first slide.

Mixed prairie grassland was a patchwork of numerous species, each of which was dominant in local microsites and associate species elsewhere. These grass species included green needlegrass (Stipa viridula), needle-and-thread (S. comata), thickspike wheatgrass (Agropyron dasystachyum), buffalograss (Buchloe dactyloides), blue grama (Bouteloua gracilis), and tall dropseed (Sporobolus asper= S. compositus) in that relative order overall. (Again, each of these grass species was a local dominant in scattered microhabitats.) Shrubs included plains pricklypear (Opuntia polycantha), silver sagebrush (Artemisia cana), and skunkbush sumac (Rhus trilobata= R. aromatica). Even taken together the cumulative cover of these shrubs did not constitute a savanna. Forbs were not common. Scarlet globemallow (Sphaeralacea coccinea) was the most widespread. Breadroot scurfpea, Indian breadroot, or prairie turnip (Psoralea esculenta) was the second-most abundant forb.

Tallgrass species were absent from this grassland such that it was a mixed prairie consisting only of shortgrass and midgrass components.

Buffalo Gap National Grassland, Pennington County, South Dakota. Mid-June, late vernal aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-59 (Wheatgrass-Needlegrass). SRM 608 (Wheatgrass-Grama-Needlegrass). Cold Temperate Grassland142- Plains Grassland 142.1, with closest being Mixed "Short-Grass" Series 142.13 in Brown et al. (1998, p. 40). Thin Upland range site (Soil Conservation Service, 1986). Northwestern Great Plains- River Breaks (of Cheyenne River) Ecoregion 43c (Bryce et al.; Undated).

8. Mixed prairie on the breaks- Two different views of Northern Mixed Prairie on flanks of a hogback in Cheyenne River breaks in the unglaciated Northern Great Plains. Major grass species (most of which are distinguishable in photographs) were green needlegrass, needle-and-thread, tall dropseed, buffalograss, and thickspike wheatgrass. Blue grama was also present, but it was too early in the warm-growing season for elongation of shoots and production of inflorescences..Tall dropseed and buffalograss were the other major warm-season (and eragrostoid) grass species. Tall dropseed was conspicuous only by presence of weathered (tattered) shoots from last year. The patch of low (short) vegetation was a colony of buffalograss (center midground of first slide; left center midground in second slide). Short, cespitose plants in left foreground of second slide were blue grama. At the right edge of this local population of blue grama were plants of plains pricklypear and small (depauperate) individuals of silver sagebrush. Plants of scarlet globemallow and prairie-turnip or breadroot scurfpea were present in this range vegetation.

This mixed prairie grassland did not have a tallgrass species and, hence, was mixed prairie of midgrass and shortgrass species.

9. Diversity of breaks grassland- Physiography of Cheyenne River breaks (including view of a hogback in right background) in the first slide and detail of physiogonomy, structure, and botanical composition of grassland (nested "photoquadrant") in the second slide. Northern Mixed Prairie of green needlegrass, needle-and-thread, tall dropseed, buffalograss, thickspike wheatgrass and blue grama with a widely spaced shrub component of skunkbush sumac (three circular-shaped shrubs in center of both photographs), silver sagebrush (behind skunkbush plants in both slides)and plains pricklypear (not visible in either of these photographs). Overall, shrubs were of such low density and cover that this range plant communiity was not a savanna (at local scale there definitely was a savanna-like physiogonomy).

On this shallow, stoney, xeric habitat the regionally dominant western wheatgrass was replaced as the wheatgrass component of mixed prairie by the drought-tolerant, disturbance-adapted thickspike wheatgrass. There was not a tallgrass species present in this range plant community. Instead, this mixed prairie consisted of midgrass and shortgrass species only. Scarlet globemallow and prairie-turnip or breadroot scurfpea, which are widespread species in the Central and Northern Great Plains, were present in small proportions (relative cover relegated these forbs to minor or even down to trace status except in very local populations).

10. Structure and composition of breaks grassland- Details of Northern Mixed Prairie on Cheyene River breaks in the Northern Great Plains at photographic distance that featured adult (and sexual) shoots of green needlegrass and needle-and-thread, two cool-season midgrass species. This was particularily prominent in the first of these two photographs. In the second photograph plains pricklypear and scarlet globemallow (a major shrub and forb, respectively) were conspicuous.

Both "photographic samples" were on flanks near the top of a hogback ridge.

11. All mixed up- Nested shots (range vegetation in second slide was closer-in view or smaller "photoquadrant" of a larger field-of-view or smaller "photoquadrant") of Northern Mixed Prairie in Cheyenne River breaks in sedimentary (unglaciated) Northern Great Plains. Grass species present in both photographs included green needlegrass, needle-and-thread, thickspike wheatgrass, tall dropseed, blue grama, and buffalograss. This mixed prairie consisted only of midgrass and shortgrass species as there was not a tallgrass species in this climax grassland.

Blue grama was conspicuous (even a pre-sexual shoot stage) in foreground of the first of these two slides. These two photographs were near enough to a major highway that there were a few plants of smooth brome (Bromus inermis) present. Smooth brome, an introduced forage species has naturalized over the Northern Great Plains Region, and its presence--even at trace cover--on this harsh, shallow-soil environment attested to its naturalized status across this vast range region. Smooth bromegrass was not, however, a major range plant in this more marginal habitat (recall that there were no native tallgrass species present).

Silver sagebrush and plains pricklypear were prominent woody species in this range vegetation. Plants of both of these shrubs were "stand-out" members of this grassland community in both of these slides.

12. 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).

13. Across the unglaciated northern plains- Landscape of Northern Mixed Prairie on the sedimentary portion of Northrn Great Plains. In the parlance of Landscape Ecology this overall (entire) grassland consisted of a matrix of western wheatgrass, blue grama, and bluebunch wheatgrass (Agropyron spictum) with patches which were co-dominated by little bluestem and (Calamovilfa longifolia). The tan- or buckskin-colored herbage (right side of second photograph) was last year's dead shoots (straw) of little bluestem which was co-dominant with prairie sandreed that grew on the outer portion and margin of these spots of little bluestem. Other locally important species included Junegrass, needle-and-thread, and threadleaf caric sedge (Carex filifolia). There were also scattered spots of thickspike wheatgrass (Agropyron dasystachyum).

This was mixed prairie with components of 1) tallgrass (made up essentially by prairie sandreed), 2) midgrass (little bluestem, western wheatgrass, bluebunch wheatgrass), and 3) shortgrass (consisting of blue grama). Forbs were extremely limited on this range with the most noticable one in this vernal community being (Achillia lanulosa= A. millefolium subsp. lanulosa). The main shrub on this grassland was silver sagebrush which was of such sparse density that physiogonomy of this prairie plant community did not even suggest a savanna-form of range vegetation. The other (and quite infrequent) shrub was soapweed or soapweed yucca (Yucca glauca) which could just as easily have been regarded as a succulent forb.

Trees in background (appearing mostly as green spots) were small groves of green ash (Fraxinus pennsylvanica). (The Greeh Ash range type was covered in Miscellaneous Forests under the heading of Woodlands and Forests.).

Dawson County, Montana. Late June, late vernal society and aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-57 (Grama-Needlegrass-Wheatgrass). SRM 606 (Wheatgrass-Bluestem-Needlegrass). Cold Temperate Grassland 142, Plains Grassland 142.1 but no appropriate series in Brown et al. (1998). Northwestern Great Plains- Montana Central Grasslands Ecoregion 43n (Woods et al, 2002).

14. Tallgrass-midgrass patch- Low hill or hummock in the unglaciated Northern Great Plains on which a little bluestem-prairie sandreed (co-dominants) community had developed. The straw (dead, tan-colored shoots) were little bluestem which tended to dominate the interior and bulk of this range vegetation while prairie sandreed dominated the perimeter and was less abundant on the inside of this plant community. Western wheatgrass and blue grama were associate species within this grassland community. There was considerable cover at local scale of Junegrass, needle-and-thread, and threadleaf caric sedge. There were also a few specimens (spots) of thickspike wheatgrass.

In this more northern portion of the Great Plains little bluestem grows as a midgrass rather than a tallgrass as it regarded to the south and east of this region. In the Northern Plains little bluestem typically has a height (and general plant size) more similar to that of western wheatgrass, sideoats grama, thickspike wheatgrass, and needle-and-thread. By comparison, prairie sandreed is a tallgrass species in the Northern Great Plains the same as in the Nebraska Sandhills. The well-represented blue grama is, of course, a shortgrass species. Thus, this form of mixed prairie contains all three general grass forms of tallgrasses, midgrasses, and shortgrasses and both sod-forming or turf grasses and bunchgrasses.

Silver sagebrush was the most common--though not abundant--shrub (silver sagebrush was more visible in the first slide of the next slide-caption set). There were also plants of soapweed yucca. This range vegetation was mixed for sure.

From the context of Landscape Ecology the little bluestem-prairie sandreed community grew as patches within the matrix of a western wheatgrass-blue grama-bluebunch wheatgrass community, the two grassland communities (matrix and patch) together forming a mixed prairie of one form or dominance type (a variant of SRM 606; Wheatgrass-Bluestem-Needlegrass) as part of the regional or zonal climax designated as Northern Mixed Prairie. For whatever reason(s) needle-and-thread was not a dominant or associate species on the plains grassland seen here (other than at local scale).

This mixed prairie range was in pristine condition. This photographer did not find Kentucky bluegrass or smooth brome in this climax vegetation, and there was but a Trace amount or perhaps "a tad more" of Japanese brome (a naturalized annual from Eurasia).

15. On the Montana plains and under the Big Sky- Details of a pristine mixed prairie comprised of two forms of grassland: 1) aggregations of or patches co-dominated by little bluestem and prairie sandreed within a larger or more extensive community of western wheatgrass, blue grama, and bluebunch wheatgrass. There was also meaningful cover of Junegrass, needle-and-thread, and threadleaf cric sedge plus spots of thickspike wheatgrass. There were some plants of silver sagebrush. The DYC (Damn Yellow Composite) the first of these two slides was lambstongue ragwort or lambstongue groundsel (Senecio integerrimus). The second slide showed little bluestem (amber-colored straw center background), prairie sandreed (right foreground), and western wheatgrass (foreground) to good advantage

Outstanding conservation; outstanding range: The following section of Northern Mixed Prairie was built from examples on two cattle ranges in the Little Missouri National Grassland in the Golden Valley of the sedimentary Northern Great Plains. Neither of these ranges had been grazed by cattle in the current growing/grazing season. Cattle dung attested to use of these two ranges the preceding (last) year. Both ranges were in Excellent range condition class--essentially pristine grassland--and stood as testimony to the fine stewardship under which these two pastures had been managed. These were examples of "how to do it right".

16. Golden grows the valley- Mosaic of true and mixed pairies--this latter a mixed prairie savanna--in Golden Valley in the unglaciated (sedimentary) Northern Great Plains. Landscape-scale view of physiography of this land form along with physiogonomy and range types of plains grassland in late spring of a very wet winter and spring. In the foreground on side of a hummock a mixed prairie of little bluestem, prairie sandreed, bluebunch wheatgrass, needle-and-thread, blue grama, threadleaf caric sedge Sandberg's bluegrass; several forbs including two species of Senecio: lambstongue ragwort or lambstongue groundsel (S. integerrimus) and grey ragwort or grey groundsel (S. canus); and the major (dominant) shrub, creeping juniper (Juniperus horizontalis), plus a few widely scattered plants of silver sagebrush and soapweed yucca. There were also some plants prairie rose (Rosa arkansana) and wild yellow buckwheat (Eriogonum flavum) which could be regarded as either subshrubs or forbs There were climax grasses of tallgrass, midgrass, and shortgrass species; forbs, and shrubs on this hummock vegetation.

This mixed prairie-creeping juniper plant community on the hummock was strictly speaking a climax savanna of tall-, mid-and shortgrass species with creeping juniper as a shrub dominant (as a climax species) along with decreaser shortgrasses and midgrasses. This range vegetation was a creeping juniper-shortgrass-midgrass-tallgrass savanna, a rather unique (or, at least, relatively restricted) climax plant community. The U.S. Forest Service interpreted and identified this range vegetation variously by such names as the little bluestem-creeping juniper habitat type or creeping juniper/little bluestem dwarf shrubland. This range plant community (unit of natural vegetation) was regarded as climax by the Forest Service (online version of Fire Effects Information System, Fire Sciences Laboratory Forest Service, Rocky Mountain Forest & Range Experiment Station). These authors noted that research and experimental-based knowledge "regarding fire effects on creeping juniper is sparse". Limited reports indicated that with most fires creeping juniper is not completely consumed (not totally topkilled), but it is a non-sprouting species which, nonetheless, regenerates by seed following fire. (As subsequent slides herein showed, creeping juniper is a prolific seed producer which is a feature common to Juniperus species.) It could be postulated (at least, speculated) that recurrent fire on this climax savanna would maintain cover (absolute or relative) of creeping juniper to more minimal proportions, and most likely at lower relative cover in comparison to the more fire-adapted grasses like little bluestem and western wheatgrass.

Grassland in the midground of this photograph was on a swale, a narrow valley or depression subject to overflow following spring snow melt or over heavy rain showers. This swale grassland community, including much of the upper slope behind it, was overwhelmingly western wheatgrass (a nearly solid-stand consociation of that species though with needle-and-thread as associate these constituting the vernal society) with blue grama as the estival dominant, and little bluestem as associate (these became the estival society). There was plains pricklypear, silver sagebrush, and fringed sagebrush as scattered shrubs in this western wheatgrass--needle-and-thread vernal society.

Another hummock of the little bluestem-creeping juniper habitat type or creeping juniper/little bluestem dwarf shrubland was in left background (in front of low butte/plateau land form). Incidentally, individual hills or small buttes in this and the related badlands land form were designated as "low knolls" by Barker and Whitman, 1989).

Little Missouri National Grassland, Golden Valley County, North Dakota. Late June, late vernal society and aspect. FRES No. 38 (Plains Grssland Ecosystem). K- 59 (Wheatgrass-Needlegrass). SRM 606 (Wheatgrass-Bluestem-Needlegrass) with swale being SRM 608 (Wheatgrass-Grama-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).

17. Golden Valley of characters- Structure and species composition of: 1) mixed prairie on a slope of a hummock or low hill (first slide) and 2) little bluestem-creeping juniper habitat type or creeping juniper/little bluestem dwarf shrubland--a mixed grass-savanna--(second slide). Range vegetation in first photograph was at the edges of three distinct local communities: 1) prairie sandreed-dominated stand with bluebunch wheatgrass, needle-and-thread, Junegrass, threadleaf caric sedge, and Sandberg's bluegrass (foreground); 2) little bluestem consociation, which was the vernal society, with some prairie sandreed, bluebunch wheatgrass, Junegrass, needle-and-thread, blue grama, and threadleaf caric sedge(left midground; dead, standing straw); and 3) western wheatgrass consociation with needle-and-thread as associate species (also some Junegrass, Sandberg's bluegrass, thickspike wheatgrass) as the vernal society and with blue grama and some little bluestem which constituted most of the estival society. There were plants of plains pricklypear, silver sagebrush, and fringed sagebrush as scattered shrubs in this western wheatgrass--needle-and-thread vernal society. The first two of these local communities were on slopes and top of a hummock while the latter had developed on a local swale or overflow range site.

The range plant community in the second slide was a creeping juniper-mixed grass savanna. Important grasses in this savanna were little bluestem (a co-dominant grass), bluebunch wheatgrass (a co-dominant grass), prairie sandreed (an associate), blue grama (an associate), needle-and-thread, Junegrass, western wheatgrass, and Sandberg's bluegrass. Thickspike wheatgrass, green needlegrass and Junegrass were minor grasses as was the naturalized Eurasian annual, Japanese brome. Threadleaf caric sedge was locally abundant and represented the grasslike plant caterory. Other shrubs or subshrubs (or, alternatively. semi-woody forbs) in this savanna were prairie rose and yellow wild buckwheat. Forbs included both grey groundsel or gray ragwort, lambstongue ragwort or lambstongue groundsel, and Lambert's crazyweed or Lambert's locoweed (Oxytropis lambertii). Creeping juniper was "loaded" (bearing a heavy crop) of fleshy seeds, a condition typical of Juniperus species (and shown in several slides below).

Range vegetation shown in both of these "photoquadrants", while different from each other in structure and relative species composition, were variants of local ecotones (transition zones at micro-scale). All these local and site-specific communities (eg. western wheatgrass consociation on the swale site) comprised a variant of the wheatgrass-bluestem-needlegrass cover type (SRM 606), the overall range plant community, it being the savanna feature/physiogonomy of some community forms that constituted the variant type.

This climax range vegetation developed in the Golden Valley area of the unglaciated Northern Great Plains.

Little Missouri National Grassland, Golden Valley County, North Dakota. Late June, late vernal society and aspect. FRES No. 38 (Plains Grssland Ecosystem). K- 59 (Wheatgrass-Needlegrass). SRM 606 (Wheatgrass-Bluestem-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).

18. Yes, this is a savanna- Range vegetation of a creeping juniper-mixed grass savanna designated variously as little bluestem-creeping juniper habitat type or creeping juniper/little bluestem dwarf shrubland (U.S. Forest Service; on line online version of Fire Effects Information System, Fire Sciences Laboratory, Rocky Mountain Forest & Range Experiment Station). In addition to this dominant shrub, the three dominant herbaceous species were little bluestem, bluebunch wheatgrass, and prairie sandreed with dominance varying locally. other important grasses included blue grama, western wheatgrass, needle-and-thread, Junegrass and Sandberg's bluegrass. Thickspike wheatgrass, junegrass, and green needlegrass were minor, native, perennial grasses. Threadleaf caric sedge, the sole representative of grasslike plants, was locally abundant. Two other shrubs or subshrubs (or, depending on interpretation, semiwoody forbs) included prairie rose and wild yellow buckwheat.The principal forbs were two Senecio species: S. canus, gray ragwort or gray . groundsel, and S. integerrimus, lambstongue ragwort or lambstongue groundsel. Other forbs included Lambert's crazyweed or Lambert's locoweed.

This climax range plant community, which developed in Golden Valley within the sedimentary portion of the Northern Great Plains, was mixed prairie ("in spades")-shrub savanna having almost unparallel diversity in structure and species composition. This potential natural vegetation included representatives of both warm- and cool-season grass species and of the three categories based on plant height and general size: tallgrass (prairie sandreed), midgrass (little bluestem, bluebunch wheatgrass, western wheatgrass, needle-and-thread, Junegrass), and shortgrass (blue grama, Sandberg's bluegrass). There grasslike plant and forb representatives as well as shrubs, providing a climax dominant for the savanna physiogonomy. Some of these plant species entered this range community as "outliers" of a western wheatgrass consociation in swale or depression habitats immediately below the hummock.

Little Missouri National Grassland, Golden Valley County, North Dakota. Late June, late vernal society and aspect. FRES No. 38 (Plains Grssland Ecosystem). K- 59 (Wheatgrass-Needlegrass). SRM 606 (Wheatgrass-Bluestem-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).

19. Short-shrub/all grass-height savanna- The creeping juniper-mixed grass savanna presented immediately above shown at close-up camera range. Two "photoplots" of the Juniperus horizontalis-tallgrass-midgrass-shortgrass savanna that developed on a hummock in the unglaciated Northern Great Plains. Joining the conspicuously short-statured creeping juniper was prairie sandreed (the dominant tallgrass); bluebunch wheatgrass and little bluestem (the dominant midgrasses); and blue grama and Sandberg's bluegrass (shortgrasses that were associate). Thickspike wheatgrass, Junegrass, and green needlegrass were native, perennial grasses of minor importance. Threadleaf caric sedge was a locally important grasslike plant. Although several species of forbs were present on this grassland (see immediately preceding captions) none were in these two "photoquadrants".

This shrub-mixed prairie savanna was remarkably diverse consisting of shrubs (though not seen in these two photographs, there were some plants of silver sagebrush as well as creeping juniper), forbs, a grasslike plant, and grasses of all three typical stature-based groups (tall-, mid-, shortgrasses). In addition there were both warm-season species such as prairie sandreed, little bluestem, and blue grama and cool-season species including bluebunch wheatgrass, western wheatgrass, Junegrass, and Sandberg's bluegrass. Furthermore, this natural range plant community was multi-layered vegetation.

Structure and species composition of major plant species was presenetd in these two slides. It was explained that even though creeping juniper is a nonsprouting conifer, it persist as a member of this climax savanna plant community because it: 1) is not usually completely topkilled by fires and 2) produces heavy yields of fleshy seeds that maintains a soil bank of reserve seeds capable of quickly regenerating this species following fire grassland fires (online version of Fire Effects Information System, Fire Sciences Laboratory, Forest Service, Rocky Mountain Forest & Range Experiment Station). This climax range plant community was named and characterized as the little bluestem-creeping juniper habitat type or creeping juniper/little bluestem dwarf shrubland (United States Geological Survey/ National Park Service, Undated; Association for Biodiversity Information).

Little Missouri National Grassland, Golden Valley County, North Dakota. Late June, late vernal society and aspect. FRES No. 38 (Plains Grssland Ecosystem). K- 59 (Wheatgrass-Needlegrass). SRM 606 (Wheatgrass-Bluestem-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).

20. Shifting emphasis- Edge of two grassland subtypes or two community components of this grassland range cover type. This was another view of the same hummock-swale location treated in the immediately preceding four slide-caption sets. In this fifth set, the downhill margin or perimeter of a creeping juniper-prairie sandreed-bluebunch wheatgrass-little bluestem savanna or dwarf scrub-grass transition that developed on a hummock (left foreground/midground) met with the uphill perimeter of a western wheatgrass consociation--as the vernal society--that developed on a swale or depression immediately below (at base of) the hummock (right foreground/ midground).

The creeping juniper-mixed grass savanna was described immediately above and the western wheatgrass consociation (again, as the vernal society) was described in the next two slide-caption sets. This interesting rangeland ecosystem was in Golden Valley.

Little Missouri National Grassland, Golden Valley County, North Dakota. Late June, late vernal society and aspect. FRES No. 38 (Plains Grssland Ecosystem). K- 59 (Wheatgrass-Needlegrass). SRM 606 (Wheatgrass-Bluestem-Needlegrass) overall, but swale was SRM 608 (Wheatgrass-Grama-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).

21. Where site and season make all the difference- A partion of bottomland habitat comprised of a swale or depression and lowest slopes of a hummock on the range landscape introduced in the five slide-caption sets above. This grassland landscape was an example of the Northern Mixed Prairie developed in the sedimentary Northern Great Plains, specifically beautiful Golden Valley. This lower-lying environment had as its vernal society two slightly different forms of climax mixed prairie: 1) a consociation of western wheatgrass with some blue grama in the swale, depression, or valley bottom and 2) a community of of western wheatgrass and needle-and-thread (dominant and associate, respectively) with blue grama and little bluestem which became dominants of the estival society on higher parts of the valley. Threadleaf caric sedge, Junegrass, Sandberg's bluegrass, and thickspike wheatgrass were present but commonly only in restricted microhabitats. Except for a few plants of western yarrow forbs were, for all practical considerations, not present. There were some plants (usually widely scattered) of plains pricklypear, fringed sage, and silver sagebrush throughout the various habitats of this lower part of the range landscape.

These two wide-angle views were of the lowland habitat or environment that was immediately below the hummock range plant community of creeping juniper-mixed grass savana treated immediately above. The second photograph was of range vegetation on the upper (slightly higher) portion of this swale, overflow, or depression habitat (near where it was contiguous with the hummock on which a creeping juniper-mixed grass savanna had developed. Needle-and-thread joined, as the associate species, western wheatgrass on this higher portion of the swale or depression.

Little Missouri National Grassland, Golden Valley County, North Dakota. Late June, late vernal society and aspect. FRES No. 38 (Plains Grassland Ecosystem). K- 59 (Wheatgrass-Needlegrass). SRM 608 (Wheatgrass-Grama-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).

22. A blue (sort of) bunch among neighbors- A plant of bluebunch wheatgrass (Agropyron spicatum) growing amid plants of creeping juniper and prairie sandreed (last year's dead, weathered leaves and part of flower-bearing tiller in right foreground). This was on a little bluestem-creeping juniper savanna in the sedimentary Northern Great Plains.

Bluebunch wheatgrass is a major forage grass on mixed prairie ranges along the western edge of the Northern Great Plains. Along this perimeter bluebunch wheatgrass is typically an associate rather than a dominant of the climax range vegetation. Notwithstanding associate status this member of the Hordeae or Tritaceae (barley or wheat tribe) is a decreaser, a member of the potential natural plant community. The value of bluebunch wheatgrass somewhat farther to the west in the eastern foothills of the Rocky Mountains and in the present day Palouse Prairie is attested to by selection of bluebunch wheatgrass as the State Grass of Montana. It has been postulated that bluebunch wheatgrass is a relict species that has persisted since the Palouse Prairie extended this far east (Heady, 1950; Weaver and Albertson, 1956, ps. 340-341).

Little Missouri River National Grassland, Golden Valley County, North Dakota. Late June; pre-boot phenological stage.

23. Typical of its species on the plains- Two plants of bluebunch wheatgrass growing on a little bluestem-creeping juniper savanna in the Northern Great Plains. Size and habit of these specimens was representative of this species in "average" and "better-than-average" years. These two plants were growing on a drier south slope so that overall size or dimensions of aboveground (shoot) portions and numbers of inflorescences were typical for harsher sites with proper grazing management. The range that these plants called home had been grazed on the light side of moderate use the preceding year.

There were some shoots remaining from the previous year, including some sexual shoots. In fact, some spikes that came through the preceding wet winter and even wetter spring still had unshattered spikelets on spikes.

Technical note: the Epson 700 Perfection (which it most certainly is not) scanner used to convert these 35mm slide images to JPEGs never could or would scan the second of these two slides correctly. Never buy anything manufactured or sold by Epson.

Little Missouri River National Grassland, Golden Valley County, North Dakota. Late June; pre-boot phenological stage.

24. 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 spicatain 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.

Two well-grounded forbs- Two composite forbs on the little bluestem-creeping juniper true prairie-savanna in the Golden Valley area of North Dakota (the sedimentary Northern Great Plains) were featured in the next two slide-caption sets. These two Senecio species provided a simple lesson in speciation in the plant community of a range ecosystem.

25. Grey in Golden Valley- Grey ragwort or grey groundsel (S. canus) growing with little bluestem (dominant), bluebunch wheatgrass and prairie sandreed (associates), blue grama, Junegrass, Sandberg's bluegrass, even some thickspike wheatgrass and green needlegrass. Co-dominant with little bluestem was creeping juniper. Forbs were quite limited.Other forbs besides the other Senecio, S. integerrimus, included Lambert's crazyweed or Lambert's locoweed.

Golden Valley County, North Dakota. Late June-peak bloom phenological stage.

26. A lambstongue in Golden Valley- Lambstongue ragwort or lambstongue groundsel (S. integerrimus) growing on a mixed grass--tall-, mid-, an d shortgrass species--creeping juniper savanna in the unglaciated Northern Great Plains. Associated species included creeping juniper and little bluestem (co-dominants), bluebunch wheatgrass and prairie sandreed (associate species) along with blue grama, Sandberg's bluegrass and minor cover of green needlegrass and thickspike wheatgrass. The other important forb, besides these two Senecio species, was Lambert's crazyweed or Lambert's locoweed.

Editorial note: the Epson 700 Perfection (which it most certainly is not) scanner used to convert these 35mm slide images to JPEGs never could or would scan the first of these three slides correctly. Never buy anything manufactured or sold by Epson.

Golden Valley County, North Dakota. Late June-peak bloom phenological stage.

27. Picture postcards of Big Sky range- A climax green needlegrass-western wheatgrass mixed prairie being used as a cow-calf pasture (white-tailed deer and pronghorn also grazed this range) in Northern Great Plains of southeast Montana. Physiography (relief or topography of the land) as well as physiography of this "mint condition" (Excellent range condition class) grassland range.

The associate range plant species on this pristine grassland was needle-and-thread which was a third native, perennial, cool-season (festucoid) grass. Needle-and-thread and green needlegrass are cespitose (tufted, bunched) grasses whereas western wheatgrass is a strongly rhizomatous, sod-forming grass.

Forbs had a minor combined cover on this mid-grss (mixed prairie) range. Forb species included silverleaf scurfpea or Indian breadroot (Psoralea argophylla); silvery lupine (Lupinus argenteus); western yarrow (Achillea lanulosa= A. millefolium), and blue, prairie, or Lewis' flax (Linum lewisii= L. perenne ssp. lewisii). There was also minor cover of the naturalized, annual, Eurasian grass, cheatgrass or downy brome (Bromus tectorum).

Big Horn County, Montana. Mid-June;late vernal aspect. FRES No. Plains Grasland Ecosstem. K-Wheatgrass-Needlegrass). SRM No. 607 (Wheatgrass-Needlegrass). Closest biotic community in Brown et al. 1998) was Cold Temperate Grassland 142, Plains Grassland 142.1, Mixed "Short-Grass" Series, 142.13, but this was wrong as should have had a "Mixed-Grass" Series. Northwestern Great Plains- Montana Central Grasslands ecoregion 43n (Woods et al., 2002).

28. Still under the Big Sky- Climax green needlegrass-western wheatgrass mixed prairie in the Northern Great Plains of southeast Montana.The first of these slides showed the physiogonomy of this range vegetation as well as the physiography (relief, topography, or terrain) of the land. The second slide served as a "photoplot" of the swar of this mid-grass grassland. In addition to the climax (decreaser) co-dominant grasses, there were some plants of needle-and-thread which was the associate species on this cow-calf range. There were four forb species well-represented on this range: silverleaf scurfpea or Indian breadroot; silvery lupine; western yarrow; and blue, prairie, or Lewis' flax. Of these four, western yarrow is on the list of 200 species in the International Intercollegiate Range Plant Identification Contest sponsored by the Society for Range Management (Stubbbendieck et al., 1992, ps. 226-227). There was some minor cover of cheatgrass or downy brome (Bromus tectorum), a naturalized Eurasian annual grass. Cheatgrass is also one of the 200 range plant species in the SRM Intercollegiate Range Plant Identification Contest (Stubbendieck et al., 1992, ps. 166-167).

It was emphasized to students that the dominant plants of this climax prairie vegetation were cool-season, perennial, decreaser grass species.

29. Big Sky Country at its base- "Photoplots" of the climax vegetation of a green needlegrass-western wheatgrass mixed prairie in the Northern Great Plains in southeast Montana. The first "photoplot" showed the climax co-dominants, the taller grass being green needlegrass, of course, while most of the shorter turf was of the rhizomatous western wheatgrass. The second "photoplot" showed the sward of this virgin vegetation that was being used as a cow-calf range. In addition to the co-dominant decreaser grasses, in this second "photoquadrant" there were plants of needle-and-thread as well as those of three forbs (including two herbaceous legumes): silverleaf scurfpea or Indian breadroot; silvery lupine; and blue, prairie, or Lewis' flax.

Note on range vegetation classification: Something of a case could be made for viewing the green needlegrass-western wheatgrass cover type (SRM 607) as a form of true prairie but a much stronger case can be presented for it being mixed prairie (See Clements, 1920, esp. 121-124, 135-138). The closeness and close proximity of true and mixed prairie in this range region was noted and this rangeland cover type remains here with a location notice given in the Grassland chapter, entitled True Prairie.

30. Representative of a commenreative species- Single plant of the perennial wild blue or Lewis' flax (Linium lewisii= L. perenne ssp. lewisii) growing on a climax mixed prairie dominated by green needlegrass and western wheatgrass with needle-and-thread as the associate species. Wild blue flax is a widely distributed species with a species range extending from Alaska eastward to Quebec and south to California back to Texas (Lesica, 2013, p. 350). Lewis' flax is not abundant enough to provide much by range feed. Just another native forb.

Big Horn County, Montana. Mid-June; final-bloom stage of phenology.

31. Only impressive up close- Upper shoot (first or upper slide) and flower (second or lower slide) of Lewis' or wild blue flax on climax mixed prairie dominated There is apparently little information regarding L. lewisii in the standard wild flower and forb literature. Hermann (1966, ps. 173, 174), Rickett (1973, p. 292), Great Plains Flora Association (1986, p. 562), and Lesica (2012, p. 350) described it briefly.

Big Horn County, Montana. Mid-June; final-bloom stage of phenology.

Northern Great Plains Mixed Prairie- Badlands

Some of the most picturesque (and certainly the roughest) range country in the Northern Great Plains is that of the various badlands that formed in the drainages of major rivers. Some of the largest and most distinctive of these are the White River and Cheyenne River Badlands of South Dakota and the Little Missouri River Badlands of North Dakota. There are also smaller areas of badlands along the Yellowstone and Missouri Rivers although these areas are often thought of, and designated more as, river breaks than badlands. Barker and Whitman (1989) described the vegetation of breaks and badlands in their exhaustive Vegetation of the Northern Great Plains. This classic work served as the basis--in more-or-less unchanged form--for descriptions of rangeland cover types of the Northern Great Plains Region in Rangeland Cover Types of the United States (Shiflet, 1994). The short section on badlands and river breaks in Barker and Whitman (1989) was omitted from the Society for Range Management publication edited by Shiflet (1994). Readers were referred to the original work.

For all of the submarginal nature of badlands and breaks range this grazing land has one indisputable feature: it cannot be farmed and will remain as range. In addition, much of this scenic (if low-production) range is now public land as National Parks and National Grasslands. The range vegetation on much of this land that "nobody wanted" is in remarkably high successional status, much of it--in spite of low production potential--is at the climax (or at least high seral) state.

The section below was devoted to badlands and river breaks range. In conjucntion with the bleak barrenness of such land are pristine grasslands, savannas, and woodlands. The author hopes that viewers enjoy it as much as Theodore Roosevelt and he did. Charge!

Little Missouri River Badlands: The Rough RiderRange Tour began with the rough country that was so central to form the extraordinary character of the Conservation President and the first wave of the Conservation Movement.

32. Short savanna just short of the badlands- At margin of Little Missouri River Badlands a rangeland community of the little bluestem-creeping juniper habitat type or creeping juniper/little bluestem dwarf shrubland. This creeping juniper-prairie grass savanna was a midgrass variant in contrast to the mixed grass variant described above. Strictly speaking this was a true prairie-savanna community or, same thing, true prairie with creeping juniper versus mixed prairie with creeping juniper treated earlier. Thus this creeping juniper-grassland (more precisely, a grass-creeping juniper savanna)--which is climax range vegetation--belonged under the heading of true prairie and, therefore, was included in the True Prairie chapter. This climax range plant community was also included here for 1) purposes of continuity (ie to have all the creeping juniper-grass savannahs together) and 2) ease of comparison with the creeping juniper-mixed grass savanna shown above (ie. for convenience of viewers).

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

48. Where lead actors change with the season- Sward of a community dominated by western wheatgrass and with needle-and-thread as secondary co-dominant or associate species on upper reaches of a swale in the Golden Valley of the Northern Great Plains. This was the point at the low area or depression joined a hummock in this range landscape. Blue grama was an associate species in this vernal society of festucoid grasses.Blue grama, which was in early stages of growth, was the dominant of the estival society. There were also some plants of threadleaf caric sedge and Sandberg's bluegrass.

Dominants of this grassland changed dramatically with advance of the growing season. A new cast of characters would appeared in the estival society of this mixed prairie as cool-season actors went into summer dormancy and fadded behind the scenes of this range in the Northern Mixed Prairie Region. Seasonal aspect and society can change drastically in the mixed prairie as in the true prairie to which the former has its gratests affinity (Clements, 1920, 138-139; Weaver and Clements, 1938, p.524), but there are fewer societies in western areas of the mixed prairie (eg. such as the mixed prairie vegetation seen here).

Little Missouri National Grassland, Golden Valley County, North Dakota. Late June, late vernal society and aspect. FRES No. 38 (Plains Grssland Ecosystem). K- 59 (Wheatgrass-Needlegrass). SRM 608 (Wheatgrass-Grama-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).

49. Another example of changing "phytoacters"- Mixed prairie range co-dominated by western wheatgrass, a cool-season midgrass, and blue grama, a warm seasson shortgrass in the sedimentary part of the Northern Great Plains. These two photographs provided a progressively shorter camera focal point to show physiogonomy, structure, and species composition of this relatively simple grassland. In some microsites needle-and-thread was locally abundant even being a cool-season associate to western wheatgrass, but in general this vernal society was a consociation of the festucoid species western wheatgrass. With advance of the yearly cycle blue grama became the dominant of the estival society which was then a consociation of this eragrostoid grass. Threadleaf caric sedge, a cool-season species, was fairly abundant in the vernal society, but blue grama (though it was in an early, prebloom stage) was the associate in the vernal society. By the way, this was mixed prairie rather than true prairie because the grassland consisted of a dominant midgrass and a dominant shortgrass species so that there was a major component of both these stature-based designations.

The "lead character" on this vegetational stage changed with the changing seasons. At such northerly latitudes in North America seasonality of the range plant community, as portrayed by climax societies, and pronounced differences in aspect are a key characteristics of the natural vegetation. Physiogonomy of this simple grassland was readily seen in these two slides.

A few isolated plants of plains pricklypear constituted the only shrubs in this mixed prairie. The only forbs found on this range by the photographer were the native (and more common or least limited) prairie onion (Allium textile) and the alien weed, leafy spurge (Euphorbia esula). These two were of such "sub-trace" proportions that they were of no practical consequence.

Little Missouri National Grassland, Golden Valley County, North Dakota. Late June, late vernal society and aspect. FRES No. 38 (Plains Grssland Ecosystem). K- 59 (Wheatgrass-Needlegrass). SRM 608 (Wheatgrass-Grama-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).

50. Simple but not boring to a grasslander- Structure and species composition of a western wheatgrass-blue grama mixed prairie range in the Golden Valley area of the sedimentary Northern Great Plains. This slide showed two forms of this amazingly simple grassland: 1) a consociation (or, perhaps, more of a colony) of western wheatgrass (left) and 2) mixture of western wheatgrass and blue grama as co-dominants (cool-season festucoid and warm-season eragrostoid, respectively). A few isolated plants of plains pricklypear added, for the record, a shrub--though not a shrub component per se--to this simple grassland climax.

The next (immediately following) slide-caption set presented a more representative composite (summary) view of this mixed prairie range.

Little Missouri National Grassland, Golden Valley County, North Dakota. Late June, late vernal society and aspect. FRES No. 38 (Plains Grssland Ecosystem). K- 59 (Wheatgrass-Needlegrass). SRM 608 (Wheatgrass-Grama-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).

51. Still not boring to a rangeman- Mixed prairie in the Golden Valley area of Northern Great Plains that was co-dominated by western wheatgrass and blue grama with a sizable proportion of the ground area "held" by threadleaf sedge. There was also respectable cover of needle-and-thread. Green needlegrass was another, though minor, member of this graminoid cast of characters. Although not visible in these photographs there were a few, widely scattered plants of plains prickly which was the only shrub present.

This pair of slides was a "nested photoplot" with the second slide being a closer-in view of the general sward seen in the first slide. The longer and larger shoots with glaucus coloration were obviously those of western wheatgrass, the cool-season co-dominant and dominant of the vernal society. Finner-stemmed and greener shoots were those of threadleaf sedge, an associate species in the vernal society, and blue grama, dominant of the estival society (and overall co-dominant of this rangeland cover type). The few greener shoots of taller height were of green needlegrass.

Little Missouri National Grassland, Golden Valley County, North Dakota. Late June, late vernal society and aspect. FRES No. 38 (Plains Grssland Ecosystem). K- 59 (Wheatgrass-Needlegrass). SRM 608 (Wheatgrass-Grama-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).

52. A tad more variety at another spot- Another photographic sample of the western wheatgrass-blue grama mixed prairie described in the immediately preceding three slide-caption sets. This pair of photographs provided another "nested photoplot" at another sampling location of this same range in the Golden Valley area of the sedimentary Northern Great Plains.

Little Missouri National Grassland, Golden Valley County, North Dakota. Late June, late vernal society and aspect. FRES No. 38 (Plains Grssland Ecosystem). K- 59 (Wheatgrass-Needlegrass). SRM 608 (Wheatgrass-Grama-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).

Dog town: The following section presented and described vegetation of severely depleted mixed prairie range on towns of black-tailed prairie dog (Cynomys ludovicianus) at three different locations in the Little Missouri River Badlands (Theodore National Park).

53. Desolate city (or a grassland this was not)- Sweeping view of part of a rather large (by present-day spatial scale) town of black-tailed prairie dog on upland, mixed prairie range in Little Missouri River Badlands. The potential natural (climax) vegetation for this grassland was western wheatgrass--little bluestem--needle-and-thread-blue grama-sideoats grama form of Northern Mixed Prairie. Through herbivory (grazing, burrowing, whatever else) of black-tailed prairie dogs this mixed prairie range had been degraded to a weed patch of wooly plantain (Plantago patagonica), field bindweed (Convolvulus arvensis), common pepperweed (Lepidium densiflorum), and yellow sweet-clover (Melilotus officinalis) as dominants (dominance of each species varying locally) with Japanese chess (Bromus japonicus) being the overall associate species. Wooly plantain and common pepperweed were the two plant species that was native to North America. The other species are natives of Eurasia that, having been brought here--purposely or accidentally--by the whiteman, naturalized, especially under conditions of periodic disturbance.

Dramatic increases in exotic grasses and forbs (mostly annual or biennial species) as shown here and in subsequent slides was precisely what Fahnestock and Detling (2001) reported for range vegetation on towns of black-tailed prairie dogs in Badlands National Park. These workers concluded that these dog towns might serve as reservoirs for seed of exotic (weedy) species. Based largely on studies conducted at Wind Cave National Park, Detlin (1998) summarized that grasses were used heavily by black-tailed prairie dogs so that grasses were put at a competitive disadvantage relative to range plants of other growth or life forms. This resulted in replacement of grasses, "often within several years". Thus, on older towns (areas colonized by prairie dogs for longer periods of time) "dominant grasses were largely replaced by forbs and dwarf shrubs" (Detlin, 1998). Coppock et al. (1983) reported that on Northern Mixed Prairie the quantity of plant biomass at peak standing crop was significantly greater (almost twice as great) on areas that were free of black-tailed prairie dogs compared to peak biomass on dog towns that had been colonized for periods of three to eight years. More important than difference in peak biomass was change in species composition of the prairie dog range. Grasses and sedges comprised more than 85% of range plant biomass on uncolonized mixed prairie, but only 5% of phytomass on prairie dog towns in which 95% of plant biomass was forbs and dwarf shrubs (Coppock et al., 1983). Fahnestock and Detling (2002) described "the near elimination of grasses on prairie dog colonies".

The only native perennial grass that this photographer could find in a quick perusal of the plant community growing on this dramatically deteriorated mixed prairie range was the shortgrass species, Sandberg's bluegrass (Poa sandbergii= P. secunda), which on this bottomland range site was an invader.

Mislabeled, misleading, misunderstood, misread, and just plain missed: Various state conservation agencies, the Nature Consrvancy, and the Association for Bioodiversity Information (Undated) in their joint publication, Plant Communities of the Midwest (North Dakota Subset)-Classification in an Ecological Context), and the United States Geologic Service/National Park Service (Undated) in their appendix, Vegetation Types and Map Units of Network Parks, described the vegetation of black-tailed prairie dog towns as: Blacktailed Prairie Dog Town Grassland Complex. Rank and scientific vegetation type of this unit was based on rank of black-tailed prairie dog unlike other vegetation types that were based on plant species.

The designation of "grassland complex" was generally flat-out wrong. The vegetation of black-tailed prairie dog towns was not grassland. Strictly speaking, the grazing disturbance climax was forbland. Furthermore it was forbland comprised to large degree by exotic noxious (=weedy) species naturalized from the continent of Eurasia following introduction by European man. Plainly and simply put, the fossorial rodents converted climax mixed prairie into a weed patch, a disclimax of alien yet naturalized forbs (annual and biennial species) with an annual grass being an associate to, infrequently, a co-dominant.

Yellow sweet-clover, a nodulated legume, is an agronomic forage (both pasture and hay) species that is a valuable range plant that is widely distributed. This aspect of Melilotis species was covered in the chapter, Introduced Forages-Legumes etc., in the Grassland group of Range Types. So, too (though to lesser degree), for field bindweed and Japanese chess. Nonetheless, it is undeniable that all these species are series pests under certain conditions and situations (Whitson et al., 1992, ps. 284, 338, 424).

Theodore Roosevelt National Park (South Unit), Billings County, North Dakota. Late June, early estival aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-59 (Wheatgrass-Needlegrass). Degraded variant of SRM 606 (Wheatgrass-Bluestem-Needlegrass) or SRM 607 (Wheatgrass-Needgrass). Blacktailed Prairie Dog Town Grassland Complex (United States Geological Survey/National Park Service, Undated; Association for Biodiversity Information). 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).

54. Front yard- Wide-scale view of part of a black-tailed prairie dog town (a colony of these fossorial rodents) on the upper terrace of Little Missouri River floodplain (first slide) and entrance of prairie dog burrow with surrounding range vegetation (second slide). This second slide was a nested "photoplot" within the larger experimental plot of the black-tailed prairie dog colony. This was the second of three colonies (towns) of black-tailed prairie dogs shown here that were in the Little Missouri River Badlands. The climax range vegetation for this bottomland (larger floodplain) site was mixed paririe consisting of western wheatgrass, needle-and-thread, prairie sandreed, Junegrass, blue grama, and green needlegrass with occasional silver sagebrush..

The present range plant community was forbland consisting of wooly plantain, field bindweed, common or prairie pepperweed, and Japanese brome. There was almost zero cover of native perennial (climax) grasses except for a few plants of Sandberg's bluegrass. There were no plants of silver sagebrush large enough to be seen above the dense foliar cover (in excess of 80%) of annual forbs and grass. Loss of western wheatgrass under heavy grazing typical of black-tailed prairie dogs on this floodplain mixed prairie site was consistent with the observation by Koford (1958) that western wheatgrass was one of the first grass species to decline in mixed prairie. Replacement of native perennial grasses from the range of prairie dog towns and replacement of these climax grasses by annual forbs and grasses was also reported for shortgrass plains range (Bonham and Lerwick, 1976).

55. Zootic (= zoological) disclimax-Sward of a grazing disturbance climax forbland comprised of wooly plantain, field bindweed, common or prairie pepperweed, and Japanese chess on the upper (highest) terrace of Little Missouri River. These two "photoquadrants" were of the range vegetation on the prairie dog town shown in the two immediately preceding photographs. There were a few plants of Sandberg's bluegrass, an invader shortgrass which was the only native species the author could find on this weed patch. The potential natural vegetation for this higher floodplain range site was Northern mixed Prairie of western wheatgrass, needled-and-thread, blue grama, prairie sandreed, Junegrass, and green needlegrass with widely scattered plants of silver sagebrush. herbivory by black-tailed prairie dogs and perhpas in conjuction with bison grazing had degraded this rangeland into this early (perhaps, pioneer) seral stage.

High proportions (relative cover, high density, general abundance) of wooly plantain and common or prairie pepperweed in this prairie dog-buffalo disclimax (a degraded mixed prairie) was consistent with findings reported for shortgrass range in the Central Great Plains where wooly plantain was frequently the dominant (had greatest cover) plant of towns of black-tailed prairie dogs (Bonham and Lerwick, 1976). Field bindweed, a major and exotic forb (noxious weed), was not found on the less mesic shortgrass range in the Central Great Plains (Bonham and Lerwick, 1976).

Weed patches (pioneer or early seral stages of vegetation; near-final stages of plant community retrogrsssion) typically persist for several years after elimination of overgrazing by black-tailed prairie dogs. Uresk and Bjugstad (1983) and Uresk (1985) concluded that in the Northern Great Plains mixed prairie ranges that were degraded to Poor range condition by black-tailed prairie dogs might require protection from all herbivore grazing for periods of four up to nine (or more) years before forage production could be increased (by extension, improved range condition class). Cid et al. (1991) found that on Northern Mixed Prairie biomass production by dominant grasses such as western wheatgrass and blue grama increased by over 30% within two years of removal of grazing by prairie dogs and/or buffalo, but that recovery of range vegetation depended on range site, extent or degree of degradation (retrogression), growing conditions, etc.

Time periods over which retrogression of mixed prairie proceeded to forbland would also with such factors. Archer et al. (1987) quantified that on Northern Mixed Prairie (inside Wind Cave National Park) annual forbs increased and dominant grasses such as buffalograss, blue grama, needle-and-thread, Kentucky bluegrass, and western wheatgrass decreased with length of colonization (time since establishment of a new town) by black-tailed prairie dogs. Within three years these perennial grasses were largely replaced by annual forbs (weeds) so that within four to six years of establishment of prairie dog towns former grassland became dominated by the annual composite, fetid marigold (Dyssodia papposa). In short there was retrogression of climax mixed prairie down to a weed patch within six years after establishment of a new town of black-tailed prairie dogs. Loss of midgrass species, decline in vegetational litter, and substantial increases in bare soil surface took place during the first two yers of new colonization by prairie dogs (Archer et al., 1987).

56. Another desolate town- The third (of a series of three) colony of black-tailed prairie dogs in the Little Missouri River Badlands. This dog town was at the highest elevation and farthest distance from the river floodplain of the three comparatively large colonies of prairie dogs presented in this section. This degraded range of a black-tailed prairie dog town was on upland slopes at the base of higher hills inside the badlands.Climax vegetation for this range environment was also Northern Mixed Prairie including little bluestem and sideoats grama (which were absent from potential or climax plant communities of floodplain prairies) in addition to western wheatgrass, needle-and-thread, blue grama, prairie sandreed, Junegrass, green needlegrass, and occasional (widely spaced) plants of silver sagebrush. Instead of such climax grassland the present range plant community of this dog town was a forbland of wooly plantain, field bindweed, and Japanese chess (ie. a weed patch).

Almost all perennial native grasses had been eliminated by herbivory (grazing, burrowing, etc,) of black-tailed prairie dogs, perhaps in combination with North American buffalo. It should be highlighted, however, that bison also grazed mixed prairie that was in climax state (= Excellent range condition class) and that was contiguous (immediately adjacent to) the degraded range of this prairie dog town on which vegetation was one of the earlier--if not first or pioneer--seral stages (a weed patch) This was a controlled experiment to the extent that there was grazing and related herbivory by 1) both bison and prairie dogs on rangeland (of the same range site) that was in direct contact with 2) rangeland grazed only by bison. (Examples of Excellent range condition class Northern Mixed Prairie of several range sites and cover types that were grazed by bison--and without prairie dogs--on grassland of this national park were shown and described above in this same section.)

57. Another prairie dog weed patch- Sward of range vegetation in a black-tailed prairie dog town on an upland, at the footslope of low hills, in Little Missouri River Badlands. This plant community of this depleted (former or once was) mixed prairie was currently a retrograde (sucessionally down the sere) community of naturalized and native plant species that were annual or biennal seral, forbs and grasses. Naturalized Eurasian species were field bindweed and Japanese chess or Japanese brome (both annuals). Native species included wooly plantain and common or prairie pepperweed. All these species were weeds as defined in either the 1) agronomic (= agricultural or economic) olr 2) ecological (= successional). There were also some plants of the domestic or introduced (from Eurasia and now naturalized) biennial, yellow sweet-clover. Herbage of all of these species (though less so for prairie pepperweed) had been utilized by black-tailed prairie dogs and buffalo (and perhaps also so by white-tailed deer and feral horses), but all these species were noxious range plants (again, weeds).

Several studies (Fahnestock and Detling, 2002) reported that native ruminants such as North American buffalo, elk, and pronghorn are attracted to range that is occupied by colonies (towns) of black-tailed prairie dogs and that these larger herbivores, ungulates (hooved mammals), are partly responsible for the overgrazed state of such grassland.

58. Combined efforts and impacts- General view of black-tailed prairie dog town with local spot where buffalo ate soil (geophagy) behind entrance to a prairie dog burrow (first slide) and close-up view of the microsite of soil consumption (second slide). Buffalo are often attracted to prairie dog towns (for whatever reasons) whereas under other conditions bison appear to avoid dog towns (for whatever reasons). Most of the plant cover and herbage in foreground of first photograph was yellow sweet-clover whereas most plant cover and biomass in background (behind earth-eating spot) was wooly plantain and field bindweed. Field bindweed is extremely palatable to cattle so it seemed likely that buffalo also found this species to their liking and were attracted to dog towns where field bindweed flourished. The author was unaware of any studies of buffalo preference for field bindweed, but his observations on various occasions proved (to him anyway) that domestic cattle readily select herbage of field bindweed. Likewise, it is common knowledge and field observation that yellow and white sweet-clovers are also palatable to numerous species of grazing animals. Buffalo and/or prairie dogs had heavily grazed (high-moderate utilization) yellow sweet-clover seen in the first photograph.

It is also likely that buffalo avoid areas of deteriorated range on dog towns that are dominated by wooly plantain. The author does not have evidence for this statement other than personal range observations and the fact that wooly plantain was generally regarded as having only fair down to poor forage value for cattle and wildlife (Stubbendieck et al.1992, p. 387). Conversely, North American buffalo might be attracted to prairie dog-degraded range on which wooly plantain is abundant if Hermann (1966, p. 271) was correct in stating that wooly plantain is "... excellent feed for all classes of stock from March to June". Studies of feed preferences and botanical contents of buffalo diets would be useful in this regard, but based on the authors long-standing experience with fistulated cattle and sheep for such studies he would caution against use of esophageal-fistulated buffalo.

Several studies and numerous field observations found that buffalo and various other native ruminants, including pronghorn (Antilocapra americana) and elk or wapiti (Cervus canadensis= C. eleaphus), are attracted to herbage (perhaps other features) on black-tailed prairie dog towns. Numerous of these studies were cited by Whicker and Detling (1988) and Fahnestock and Detling (2006), the former duo of which summarized previous work and the latter of which reported on nutrient cycling and nitrogen content of plant shoots on towns of black-tailed prairie dogs. Concentrationss of nitrogen and/or crude protein of plants on black-tailed prairie dog towns were higher than on control plots with no prairie dogs on Northern Mixed Prairie in both Wind Cave Nationl Park (Coppock et al., 1983b) and Badlands National Park (Fahnestock and Detling, 2002). Buffalo appeared to be attracted and graze by preference in areas colonized by black-tailed prairie dogs ("dog towns") as a result of more nutritious forage as well as different vegetational structure which probably included the feature of lower plant stature (Coppock et al., 1983b; Whicker and Detling, 1988).

This degraded range of a black-tailed prairie dog town of was on upland rangeland inside the Little Missouri River Badlands. This was a different general range environment from the first two examples of black-tailed prairie dog towns which were on the outer (higher) terraces of the Little Missouri River floodplain (see above).

Concluding (though tentative) remark: the published literature regarding impact of the black-tailed prairie dog on range vegetation is conflicting, contradictory, and contentious, and in at least one isolated case, even contemptible (as in fradulent). Most importantly, perhaps, is the fact that it is extremely limited in comparison to other topics related to black-tailed prairie dogs. Studies of black-tailed prairie dogs appearing in peer-reviewed journals included frontier travelers' accounts, natural history, economic analyses, geographic distribution of the species, dynamics of populations, behavior including behavioral ecology, associations and interactions with other animals, and many others, but amazingly there was very little science as to successional effects (if any) of these dominant native animals on range vegetation. For example, in a grazing trial designed to determine if (and if so, how much) cattle performance was affected by black-tailed prairie dog herbivory (Derner et al., 2006) composition and production of range vegetation was not measured.

What is more (and this is strictly "secret talk"), at least one of the more prominent peer-reviewed papers that claimed to have examined range vegetation on black-tailed prairie dog towns and compared these plant communities to those on control plots was a fraud with the hidden political agenda to show that black-tailed prairie dogs had minimal impact of range vegetation. In what appeared to the present author as an even-handed treatment of the subject of prairie dog management, Stapp (1998) concluded that as native rodents that were integral parts of natural grasslands performing critical roles in these range ecosystems prairie dogs merited conservation. He also concluded: "... it is unfortunate that there seems to be a perceived need for selective representation of published studies to create a greater ecological role for prairie dogs than is justified by existing data" (Stapp, 1998).

Nonetheless, there have been a number of solid, exemplary studies on impacts of black-tailed prairie dogs on range vegetation. A number of these were done by graduate students at Colorado State University with findings published in theses. These were listed by Hyde (1981) and at least one study (Bonham and Lerwick, 1976) on vegetation (shortgrass range) changes was reported. Other sound, relevant studies included those of Koford (1953), Coppock et al (1983a, 1983b), Uresk (1985), Archer et al (1987), Weltzin et al. (1997), Detling (1998), Fahnestock and Detling (2001), Derner et al. (2006). Whicker and Detling (1988) and Detling (1998) furnished summaries of black-tailed prairie dog impacts of grasslands complete with extensive literature citations.

As this discussion did not purport to be a literature review the author left it up to readers to search on their own through the comparatively vast volume of peer-reviewed literature regarding any of the black-tailed prairie dog-related aspects listed in the first paragraph of this concluding remark. A large number of such investigations can be located by searching through the Literature Cited of studies listed above. Published in scientific periodicals or not, the blaringly, immediately obvious fact to any astute observer is that the range plant community within a black-tailed prairie dog town differs drastically from that of adjacent grazed range. Nowhere is this more striking than on public land administered by federal conservation agencies. The examples presented above from Theodore Roosevelt National Park were a case in point. This author witnessed this same situation on mixed prairie in Wind Cave National Park and on tallgrass prairie in the Nebraska Sandhills on Fort Niobrara National Wildlife Refuge. The latter was presented and discussed herein under the chapter, Tallgrass Prairie. It should be emphasized that in all three of these cases the other dominant range herbivore was North American buffalo. The range was not being grazed by livestock at time of photographs and accompanying descriptions of range plant communities in Range Types of North America. Another example of range deterioration by black-tailed prairie dogs on shortgrass plains range that was grazed exclusively by prairie dogs was presented above under the Central Great Plains section.

This author would underscore one fact: much, prehaps most, of the herbaceous cover and biomass on black-tailed prairie dog towns shown immediately above was of Eurasian species introduced by European man. In absence of these exotic, largely weedy plants the vegetation on the ranges presented above could have been different (ie. presumedly native plant species would have made up the range vegetation in absence of exotic opportunistic weeds or, in case of yellow sweet-clover, a naturalized domestic forage crop. On the other hand, there were areas on prairie dog-degraded rangeland that supported populations and/or simple communities comprised of native annual forbs (eg. wooly plantain, common pepperweed) that are pioneers in the first seral stage of secondary succession. These species are the colonizers of abandoned farmland indicating the dramatic state of retrogression on black-tailed prairie towns such as those seen above.

In the same vein, diminished numbers or total absence of native predators such as black-footed ferret (Mustela nigripes) and, conversely, introduction of exotic diseases to which the prairie dog has not evolved resistance such as Bubonic Plauge caused by the bacteium, Yersinia pestis, in all probably had some impact on the black-tailed prairie dog populations that could have influenced its impacts on grasslands.

The black-tailed prairie dog has been interpreted as a keystone species on the central grasslands of North America (Fahnestock and Detling, 2002). Stapp (1998) felt that "it may be premature" to assign keystone species status to prairie dogs pointing out that their impact varied greatly between shortgrass plains grasslands where there were less drastic changes in species composition than on mixed prairie grasslands. Nonetheless, Stapp (1998) concluded that prairie dogs were important components of grassland ecosystems and landscapes, critical for presence of other vertebrates, and worthy of protection from wholesale extermination projects such as those of the past. Uresk and Bjugstad (1983) labeled black-tailed prairie dogs as "ecosystem regulators". Whicker and Detling (1988) described the black-tailed prairie dog as having a "natural role as agents of intense vegetational disturbance" and then they quite logically and cogently described the impact of these comparatively large rodents in the context of creating patches in the matrix of mixed prairie with vegetational dynamics that followed the general processes of plant retrogression/succession in the pattern of what has become known as patch dynamics (Pickett and White, 1985).

In this publication, the author deemed it sufficient to show changes in range vegetation on towns of this native, fossorial rodent. Readers were left to draw their own tentative conclusions from whatever knowledge could be gleaned from this presentation. Given the state of the scientific literature--at time of this writing--it did not seem that the peer-reviewed literature offered any more readily obvious evidence that what was obtained and shown by nothing more advanced than color photography.

White River Badlands: In some respects the White River Badlands appear even more stark, remote, forlourn, and forsaken than those of the Little Missouri River. This perception is perhaps enhanced due to proximity of the former to the Black Hills. C.C. O'Hara (1920, p. 19) stated the case best in in the seminal and forever classic work, The White River Badlands: "The White River Badlands constitute the most important badland area in the world". It has been remarked on various occasions that based on climate, especially precipitation, and vegetation it would have been more rational to have had East Dakota and West Dakota instead of North Dakota and South Dakota.. Of course, professor C. Etzel Pearcy, the California geography professor who proposed a 38-state Union, envisioned more-or-less reuniting the two Dakotas as one state of Dakota. Dream on Etzel. On to the White River Badlands.

The badlands associated with the greater White River Valley are the remains of a former valley that is being geologically eroded into a plain. Due to variation in parent materials, slopes, aspects, etc. there is a range in degrees of land erosion (differential erosion rates) with a corresponding array of ages of soils, heights of hills, and general extension of the badlands landform. Thus there are pinnacles, spires, knobs, columns, and land masses that are nothing but small remains of badlands scarcely larger than an ant mound or a mole hill. Some of these various micro-land forms may be bare (like an ant hill) or, in other cases, at some stage of plant succession. Generally, however, there will be some climax vegetation on all intermediate stages of geologic erosion (land and soil formation). In other words, the range plant community on most badlands land forms will be about or, at least closely approaching, that of the ultimate expression or development of potential natural vegetation (ie. it will be the same, or nearly the same, as the climatic or regional climax on all sizes of eroded land--local microland forms--of the badlands). The obvious exceptions are raw or freshly eroded badlands. Obviously, there are a number of range sites within the White River Badlands.

Those familiar with the scheme of vegetation development based on the geologic/climatic cycle-based proposed in Plant Succession (Cements, 1916) can visualize changes in plant communities over course of Earth history in what are now the White River Badlands. Though dated (and, to most current vegetation scientists, arcane) the conceptual interpretations by Clements (1916) can help students of vegetation understand the dynamic nature of plant communities over geologic time on land where active earth-changing processes are most apparent. Interested readers were directed to Clements (1916, especially chapters XII through XIV [ps. 279-422]). Earlier in Plant Succession Clements (!916, p. 183) presented his outline of vegetational units over course of geologic time and climate changes associated with this temporal aspect.

Land surfaces resulting from geologic erosion in the badlands constitute or comprise a temporal scheme collapsing in shorter time spans of eosere to clisere down to biosere (or, simply, sere for short) The temporal scale of sere occurs at a place where (and when) plant succession--plant community development--roceeds faster than does geologic erosion. Plant community changes from earliest stages to terminal state, the climax, is the process of vegetation development known as succession. This proceedes along paths with stages (communities) of vegetation development in a series of cycles within ever longer cycles extending back over course of geologic time: (eoseres comprised of shorter-time cliseres in turn made up of bioseres or, more simply, seres. Restated, there are a series of bioseres or, simply, seres on each clisere or eosere: there are cycles or complete sequences of vegetational development or seres. A number of seres (= bioseres), each with successional stages from pioneer to climax, comprise a clisere. Evidence of cycles of change within ever-longer cycles of change could be found in present vegetation. For example Clements (1916, ps.114-116) explained how zones in river valleys presented evidence of changes in climate and, thus, changes in climaxes that condensed time over relatively short distances.

Development of range vegetation in the White River Badlands can be viewed as both chronosequences and toposequences: plant succession (both on-going and prehistoric) took (is taking) place both on the same sere (chronosequence, a chronological or time-based sequence on the same location) and on different seres (toposequence, a topographic or location-based sequence at the same point in time). Plant succession in badlands is predominately xerarch, plant community development on dry environments. On the other hand, some development of range vegetation (some plant succession) in the White River Badlands is hydrarch, community development on wet environments. Examples of both xerarch and hydrarch forms of range plant succession were presented in this section. Plant succession on the harsh habitats of the White River Badlands can be primary succession on "new land" created by geologic erosion or, alternatively, development of range plant communities can be redevelopment of pre-existing plant vegetation via secondary succession (re-vegetation) as recovery from drought, farming, overgrazing, roadbuilding, and so forth. On the steep surfaces of the badlands land form gravity is a major agent of erosion that works in unison with water and wind in carving the stark and erie shapes and sizes of this on-going creation. Land formation and soil genesis--in conjunction with vegetation development--take place even as rangemen and other students of Nature look on in amazement and wonder.

Geologic formations including layers of fossil soils, one overlying another, along with treasure troves of plant and animal fossils in the current White River Badlands provide palaeontological evidence of these successional cycles within longer successional cycles. A total of 87 fossil soils have been recorded in Badlands National Park. Approximately 80 million years Before Present (BP) the area of present-day White River Badlands was an inland sea. Pierre Shale is the geologic stratum created out of the mud of this prehistoric body of water. Then about 65 million years BP earth-modifying forces raised the former seabed that eventually became a jungle. From the early Oligocene through the Eocene (roughly 57 to 26 million years BP) the land was changed drastically (though ever so slowly). The Laramide Orogeny (mountain-building processes that formed the laramie Range) produced the Black Hills which as they rose their streams buried the previous ocean bed that was turned into river deltas that became swampland during the Oligocene Epoch.

As the land continued to rise, the streams (some as large as rivers like White River) moved faster so as to erode away rather than cover or bury the former swamp. This geologic process created the White River Badlands Wall between the upper (higher) grasslands to the north and the lower grasslands to the south. Today this formation is most prominent at Cedar Pass. Today this is a watershed with the upper grassland part of the catchment draining into the Cheyenne and Missouri Rivers while the lower grassland of the divide drains into White River. These three rivers are tributaries into the Missouri River, and thence into the mighty Mississippi River (Froiland, 1990, ps. 177-191).

Natural or geologic erosion has resulted in deeply dissected uplands where weakly consolidated sediments have been cut away leaving the more erosion-resistant sediments. All these sediments were laid down during the Cretaceous Period (during which Pierre Shale was formed) all the way through the Eocene (when stream deposition resulted in the Chadron formation including claystone, limestone, and even channel sandstone). In the Oligocene the Brule Formation was formed that included both limestone and sandstone.

There is considerable variability or diversity in soils of the White River Badlands and immediately adjacent areas. Soil series--to the extent possible--were listed in conjunction with range sites in this section. In general, many of these soils are saline, alkaline, calcareous, often residual, frequently alluvial, deep to shallow, primarily well-drained, and with well-developed profiles.

On-going cycles of range vegetation development occur in the White River Badlands where there are all sizes (both base and height) and shapes of badlands land forms resulting from differential erosion (varying rates of geologic erosion) as a consequence of parent and other geologic materials of various hardnesses and resistances to weathering. These local badlands land forms vary from towering wall to "badger den"-sized humps with all manner of intermediate stages and sizes. Vegetation, including plant succession in particular, in the White River Badlands has not been studied as thoroughly as has general geology. Likewise, the fossil record of plants is not as rich as that of animals.

Perhaps the best overall treatment of the White River Badlands, especially from the perspective of Paleozoology, remains that of O'Hara (1920, and still in print). Also highly recommended (and one of the major sources for the above brief review) was Hauk (1969). For more recent and greater detailed coverage on geology of the White River Badlands refer to Stoffer (2003).

Barker and Whitman (1989) gave the following introductory sentence in their description of the Badlands and River Breaks unit in Vegetation of the Northern Great Plains: "The vegetation of the badlands and steep river breaks throughout most of the Northern Great Plains is so distinctive and unique that it deserves separate classification". Unfortunately--and unlike other units of vegetation in Barker and Whitman (1989)--range plant communities of badlands and breaks was not included included in Rangeland Copver Types of the United States (Shiflet, 1994). As such, there were not appropriate SRM units for grassland vegetation of White River Badlands.

59. Panarama of Mako sica (Sioux) or Les Manvaises Terres a Traverse (French)- Two languages; same meaning. The Sioux term meant "land, bad" whereas the French phrase said ""bad lands to travel across". It was left to American explorers, geologists, geographers, and settlers to give this terrain or land form a colorful name, and one based on the main land-forming agent: White River Badlands. These two panaramic views presented the kialeidoscope landscape of buttes, mesas, towers, ravines, canyons, etc. at the mature stage of erosional dissection. This was a finely textured drainage pattern with steep slopes due to differential erosion. The finer clays eroded raster and more quickly than beds of sandstone or ilsolated concretions which are harder and, therefore, erode slower.

This image of the White River Badlands were photographed looking south toward White River and the lower grasslands (referred to above). Clearly there was almost no plant life on the most severely eroded surfaces. Limited range vegetation in immediate foregrounds consisted mostly of rubber rabbitbrush (Chrysothamnus nauseosus) and western wheatgrass on upper margins and edges of eroded land surfaces.

Badlands National Park, Jackson County, South Dakota. July. Badland Map Unit (Soil Conservation Service, 1987). White River Badlands Ecoregion- 43h (Bryce et al., 1996).

60. More of the kaleidoscopic panarama- Spectacular terrain of White River Badlands taken from various overlooks along Loop Road of Badlands National Park. Obviously there was little or no vegetation on the most severely eroded faces of this land that varied in shape or form from canyons and ravines to towers and pinnacles with spires, buttes, mesas, etc. between. At bases and gentler slopes of these often cliff-like land shapes there was grassland and shrub-grass savanna vegetation much of which, when compared to surrounding mixed prairie, was apparently at the climax state.

Range vegetation (primarily grassland of the mixed prairie type) on bottoms, benches, and more advanced erosional surfaces of the White River Badlands as well as that of rangeland immediately surrounding the badlands was presented immediately below. In the photographs presented here (and immediately above) emphasis was laid on the badlands land form per se. These sweeping composite views presented all different stages of geologic erosion (and some development of vegetation). Varying degrees of hardness of geologic materials (including parent materials) and soils (both present-day and fossil soils) are the primarily explanation of the unique diversity and stupendous shapes and sizes in land surfaces. Geologic materials include mudstone, siltstone, and shale in the Badland mapping unit (Soil Conservation Service, 1988). Weakly cemented parent materials results in high drainage density (= fine texture of drainage pattern) of these badlands. There is anywhere from 30 to 200 and, sometimes, up to 500 channel miles per sqaure mile of land area.

Much of the surface area shown here was the Brule geologic formation which rests atop the Chadron formation both of which are of the Oligocene Epoch. Rapid rates of surface runoff from the upper portions of the White River Badlands result in copious quantities of water available for plants growing below and development of range vegetation.

Zones or bands of red and yellow coloration on eroded slopes and faces are fossil soils. (At least these were reds and yellows captured as only Kodachrome could "catch" them, but by the time the Epson Perfection 700 scanner ["perfection" it is not] these colors could wind up being anything but the original color.).

Badlands National Park, Jackson and Pennington Counties, South Dakota. July. Badland Map Unit (Soil Conservation Service, 1987). White River Badlands Ecoregion- 43h (Bryce et al., 1996).

61. A bad wall and its shelf- White River Badlands Wall, the remnant of a valley formed from the Laramide Orogeny , along with its cliff shelf. This geologic remnant presently separates the terrain of the upper grassland from that of the lower grassland towering 450 feet above the latter and 150 feet above the former grassland. The upper grasslands are part of the watershed of Bad River, a tributary of the Missouri River, while runoff from the lower grasslands drains into another tributary of the Missouri, white River. A large earthen slump known as Cliff Shelf, was presented in this midground and foreground of this slide. A easily read description (and a general enjoyable read) of Cliff Shelf and the Wall--along with many other aspects of the White River Badlands--remains the classic Badlands- its Life and Landscape by ranchwoman Joy V.Hauk (1969).

Natural vegetation on this part of Cliff Shelf (on the lower grassland) varied from juniper woodland (see next slide) to wet prairie. This juniper woodland was the feature of the immediately succeeding slide. The wet or ponded grassland was described immediately thereafter.

Cedar Pass, Badlands National Park, Jackson County, South Dakota. July. White River Badlands Ecoregion- 43h (Bryce et al., 1996).

62. Wood below the wall- On the highest vegetated part of Cliff Shelf, on the lower grasslands below White River Badlands Wall, a woodland had developed that was dominated by two Juniperus species: 1) Rocky Mountain juniper (J. scopulorum) and 2) eastern red cedar (J. virginiana). These two species (respectively, left and right) were shown in this slide. The nearly dormant (drought dormancy on this xeric site) herbaceous understorey consisted primarily of blue grama, buffalograss, and local patches of western wheatgrass. In some places these two Juniperus species formed a closed-canopy forest of short-statured trees with little or no herbaceous layer. This latter situation could have resulted from fire exclusion/fire suppression practices.

Juniper woodland was limited generally to the upper and drier zones of the White River Badlands, but as was shown in the next two photographs, young junipers had established and appeared to be doing well on lower and wetter parts of Cliff Shelf.

Cedar Pass, Badlands National Park, Jackson County, South Dakota. July. FRES No. 35 (Pinyon-Juniper Shrubland Ecosystem). K-21 (Juniper-Pinyon Woodland). SAF 46 (variant of Eastern Red Cedar with "recognizable percentage of Rocky Mountain juniper"). White River Badlands Ecoregion- 43h (Bryce et al., 1996).

63. What a mixture (and one that defies classification)!- Biologically diverse range plant community (communities) that developed on wetland habitats at base of Cliff Shelf and Badlands Wall in White River Badlands. Abundant soil water and shallow surface aquifers afforded this example of hydrarch plant succession, the development of range vegetation on a mesic to hydric environment. At the base of this badlands cliff there was abundant water for more favorable plant establishment, growth, and reproduction than in adjacent and surrounding habitats. This local abundance of water was due to a combination of factors including tremendous quantities of water flowing downslope (abundant runoff), throughflow (lateral flow) of water through soil, and higher water-holding capacity of clay particles in the soil.

Tree species included eastern cottonwood (Populus deltoides subsp. monilifera), eastern red cedar, and Rocky Mountain juniper. (An example of juniper woodland on Cliff Shelf was treated immediately above.) Dominant grasses were switchgrass and western wheatgrass with needle-and-thread, green needlegrass, and prairie cordgrass the associate grass species. Canada wildrye was also present. Squirreltail bottlebrush (Sitanion hystrix) formed local colonies. The major grasslike plants were broadleaf cattail (Typha latifolia), various caric sedges (Carex spp.), and an unidentifiable bulrush (Scirpus sp.) . Rubber rabbitbrush and skunkbush sumac (Rhus aromatica= R. trilobata) were the two principal shrubs. Western snowberry (Symphoricarpos occidentalis) was the associate shrub.

Viewers should note that seedlings and saplings of Juniperus species had invaded the meadow-like (herbaceous) portions of this wetland. A return to the natural fire regime (after nearly a century of fire suppression) might be expected to prevent establishment of trees of adult or advanced-juvenile stages. This would more likely be the case for the two non-sprouting Juniperus species.

Classification of this wetland vegetation was beyond this author's pay scale. This range vegetation could be viewed as two (herbaceous and woody) or more plant communities. The wet--at least, seasonally wet--meadow (the distinct herbaceous zone or zones) were wet prairie or, perhaps, marsh. The shrub-dominated zone(s) appeared to be transitional to the tree-dominated vegetation. This latter might have been a transition between juniper woodland (see immediately above) and a form of eastern cottonwood gallary forest (minus a stream). Alternatively, the tree-dominated community might be interpreted as a mesic or hydric variant of juniper woodland. All of this range vegetation was part of the general Northern Mixed Prairie Region. Some of the possible vegetation classification units were shown next.

Cedar Pass, Badlands National Park, Jackson County, South Dakota. July. Tree-dominated vegetation: FRES No. 35 (Pinyon-Juniper Shrubland Ecosystem), K-21 (Juniper-Pinyon Woodland), SAF 46 (variant of Eastern Red Cedar) or, alternatively, FRES No. 17 (Elm-Ash-Cottonwood); No fitting Kuchler unit; SAF 235 (variant of Cottonwood-Willow); Plains and Great Basin riparian Deciduous Forest 222.2, Cottonwood-Willow [minus willow] Series 222.21 of Brown et al. (1998, p. 43)with "recognizable percentage of Rocky Mountain juniper"). Herbaceous-shrub vegetation: FRES No. 41 (Wet Grassland); K-65 (Northern Cordgrass Prairie); no appropriate SRM unit for northern plains wet prairie; Cold Temperate Marshland 242, Plains Interior Marshland 242.3, No appropriate series in Brown et al. 9998, p. 45). Sub-irrigated range site. White River Badlands Ecoregion- 43h (Bryce et al., 1996).

64. Good overflow in badlands- A grassland community of prairie sandreed, switchgrass, and western wheatgrass growing as a mosaic of local consociations (each of the three grasses grew as single-species stands or populations) that had developed on a Badlands Overflow range site. This was interpreted as a climax mixed prairie with western wheatgrass, a midgrass, regarded as the potential dominant decreaser on shallower clay microsites while the two tallgrasses were dominant decreasers on microsites of deeper soils with somewhat less clay.

At this time in early summer the cool-season grasses and threadleaf sedge were mostly in the dormant stage. The typically conspicuous needle-and-thread had long-since shed its grains. Western wheatgrass was at or nearing the grain-shatter stage. Prairie sandreed and switchgrass shoots were at the boot stage on this site.

Soil in most of foreground was the Interior series as a loam or channeled loam (Soil Conservation Service, 1987).

In these two views there were actually two distinctive grassland communities (two range sites) deep in the interior of White River Badlands. A Silty range site was in the distant background. Soil of the Silty range site was Cedar Pass silt loam. The grassland of the Silty range site was featured in the two slide-caption set immediately below.

On both of these range sites (and grassland communities) the rate of overland flow from higher (and sparsely vegetated) slopes was so great (high) that the mixed prairie vegetation was postclimax, more mesic than the regional or zonal climax ("climatic" climax) of this part of the semiarid Northern Great Plains. Nonetheless, both of these range plant communities were part of the general Northern Mixed Prairie.

Badlands National Park, Pennington County, South Dakota. July. FRES No. 38 (Plains Grasslands Ecosystem). K-61 (Wheatgrass-Grama-Buffalograss) was generic at type (versus range site) scale for White River Badlands. Cold Temperate Grassland 142, Plains Grassland 142.1, Mixed "Short-Grass" Series 142.13 was closest thing--and that missed it--in Brown et al. (1998, p. 40). No appropriate SRM unit for this badlands mixed prairie: the badlands grassland mosaic described (albeit briefly) in Barker and Whitman (1989) was not included in Society for Range Management cover types (Shiflet, 1994). Badlands Overflow range site (Soil Conservation Service, 1987). White River Badlands Ecoregion- 43h (Bryce et al., 1996).

65. Still in the badlands?- Yep. Deep inside the White River Badlands a Silty range site supported a climax (a a textbook example of) mixed prairie that was essentially a consociation of western wheatgrass with other species being green needlegrass, needle-and-thread, buffalograss, sideoats grama, blue grama, hairy grama, and threadleaf sedge. There was not a tallgrass component in this midgrass-shortgrass mixed prairie. It was possible that the tallgrass species such as prairie sandreed or, perhaps more likely, little bluestem had been severely depleted by overgrazing or, even, attempted farming in the distant past. (Obviously this rangeland was not being overgrazed currently.) Contrariwise, it was possible that the central part of this large area of grassland was far enough away from higher ground that it did not receive adequate runoff (= overland flow) to support tallgrasses in this semiarid precipitation zone.

The consociation of western wheatgrass was most prominent in foreground in the first of these two slides. There was also a large local stand or colony of western wheatgrass accompanied by heavy cover of needle-andthread in center of the second slide. Most of the range vegetation in foreground of this second slide was threadleaf sedge with a lower layer (not visible) of buffalograss. The most conspicuous forb was annual sunflower (Helainthus annuus) that was present in the first slide as isolated plants. Both needlegrasses (Stipa spp.) were past grain-shatter stage. Western wheatgrass was at, or nearing, grain-ripe stage of phenology. Buffalograss had produced some burs, but seemed ready to bloom again.

Soil in the first slide was primarily Norka sild loam with small areas of Blackpipe Clay loam. Soil in background of this first photograph was Fairburn clay loam that served as basis of a Thin Upland range site. Soil in the second slide was mostly Cedar Pass silty clay loam, level.

Badlands National Park (first slide) and Buffalo Gapp National Grassland (second slide), Pennington County, South Dakota. July. FRES No. 38 (Plains Grasslands Ecosystem). K-61 (Wheatgrass-Grama-Buffalograss) was generic at type (versus range site) scale for White River Badlands. Cold Temperate Grassland 142, Plains Grassland 142.1, Mixed "Short-Grass" Series 142.13 was closest thing--and that missed it--in Brown et al. (1998, p. 40). No appropriate SRM unit for this badlands mixed prairie: the badlands grassland mosaic described (albeit briefly) in Barker and Whitman (1989) was not included in Society for Range Management cover types (Shiflet, 1994). Badlands Overflow range site (Soil Conservation Service, 1987). White River Badlands Ecoregion- 43h (Bryce et al., 1996).

66. Where the clay settled- Two examples of remarkably diverse mixed prairie that developed on heavy clay soils in White River Badlands. Grassland vegetation in the first photograph was comprised of tallgrass species (primarily prairie sandreed and switchgrass with some big bluestem), midgrasss species (western wheatgrass, green needlegrass, needle-and-thread with lesser amounts of sideoatsgrama and little bluestem), and shortgrass species (blue grama as dominant and buffalograss as associate shortgrasses). Two noteworth points: 1) the major midgrasses were cool-season species whereas all tallgrasses and the predominant shortgrass species were warm-season species and 2) little bluestem was more of a midgrass on badlands range.

Grassland vegetation in the second photograph was mixed prairie of midgrasses (a consociation of western wheatgrass) and tallgrasses (prairie sandreed and switchgrass almost exclusively) without a shortgrass component. There were a few scattered plants of blue grama and fewer yet of threadleaf sedge and buffalograss, but taken together these did not comprise a vegetational layer. This range plant community is one of numerous forms of Northern Mixed Prairie.

Both range plant communities had developed on a Clayey range site, but the two Clayey sites were of different soils. The grassland plant community of the first slide was a mixture of Cactusflat (series) silty clay and Cedarpass-Interior-Denby (three series) Complex. Clay contents of Cedarpass, Interior, and Denby series were 18 to 30%, 18 to 35%, and less than 35%, respectively. The grassland vegetation in the second slide was on Orella-Badlands Complex mapping unit (all shallow clay soils). Clay contents of the Orella series ranges from 35 to 70% clay. A Badlands Overflow range site was in background of this second slide. Examples of the Badlands Overflow range site (in a mosaic with several range sites) was presented below.

Badlands National Park, Pennington County, South Dakota. July. FRES No. 38 (Plains Grasslands Ecosystem). K-61 (Wheatgrass-Grama-Buffalograss) was generic at type (versus range site) scale for White River Badlands. Cold Temperate Grassland 142, Plains Grassland 142.1, Mixed "Short-Grass" Series 142.13 was closest thing--and that missed it--in Brown et al. (1998, p. 40). No appropriate SRM unit for this badlands mixed prairie: the badlands grassland mosaic described (albeit briefly) in Barker and Whitman (1989) was not included in Society for Range Management cover types (Shiflet, 1994). Clayey range site (Soil Conservation Service, 1987). White River Badlands Ecoregion- 43h (Bryce et al., 1996).

67. More densely settled (and less diverse)- Mixed prairie on a heavy clay soil in White River Badlands. Grassland vegetation on this cattle (cows and calves) range was a consociation of western wheatgrass with green needlegrass as the associate species. This rangeland plant community had developed on a Dense Clay range site (in contrast with or comparison to the two examples of a Clayey range site presented in the immediately preceding two slide-caption set). Another example of the diversity of native vegetation of the Northern Mixed Prairie Region.

The predominant soil series of this Dense Clay range site was Whitewater Clay that has clay contents ranging from 40 to 60% clay.

Buffalo Gap National Grassland, Pennington County, South Dakota. July. FRES No. 38 (Plains Grasslands Ecosystem). K-61 (Wheatgrass-Grama-Buffalograss) was generic at type (versus range site) scale for White River Badlands. Cold Temperate Grassland 142, Plains Grassland 142.1, Mixed "Short-Grass" Series 142.13 was closest thing--and that missed it--in Brown et al. (1998, p. 40). No appropriate SRM unit for this badlands mixed prairie: the badlands grassland mosaic described (albeit briefly) in Barker and Whitman (1989) was not included in Society for Range Management cover types (Shiflet, 1994). Dense Clay range site (Soil Conservation Service, 1987). White River Badlands Ecoregion- 43h (Bryce et al., 1996).

68. Where it settled from- Botanically rich mixed prairie cattle range in White River Badlands. This grassland plant community was a diverse complex of the following species (in this relative or more-or-less order of cover); green needlegrass, western wheatgrass, needle-and-thread, blue grama, little bluestem, sideoats grama, and threadleaf caric sedge. Tallgrass species were absent (at least essentially so) from this rangeland vegetation that was primarily on a Thin Upland range site with patches of Shallow Clay range site.

Remnants of the former bed of the Chicago, Milwaukee, St Paul and Pacific Railroad, the Milwaukee Road, were in the midground of this interesting view of White River Badlands grassland. Part of the Badlands Wall was visible against the distant skyline. What stories this land ancient land could tell. An ever-unfolding geologic stage of inland seas and, later, swamps in the Age of the Mammals to the current mixed prairie association of the great North American interior grasslands. A tale of "God's Country" from the Late Cretaceous through Oligicene and Miocene Epochs dating from almost 80 million years ago (see Figure 9 in Hauk, 1969). Earliest human inhabitants were the Asian immigrants, the Paleo-Indians who arrived via the Berring Land Bridge. Later, the Arikara people laid claim to this part of the Western Range. The Arikara were followed by the Lakota who later merged with the Western Dakota and Eastern Dakota to form the mighty Sioux Nation. Finally, the white man and his iron horse such as the once-mighty Milwaukee Road. The tale continued with the bitter-sweet story of annihiliation of the buffalo (Bison bison) and the red man. The pagent continued to the present era of wise use and preservation of these natural resources for those of future generations who can continue to marvel at the wonders of this ancient land.

Buffalo Gap National Grassland, Pennington County, South Dakota. July. FRES No. 38 (Plains Grasslands Ecosystem). K-61 (Wheatgrass-Grama-Buffalograss) was generic at type (versus range site) scale for White River Badlands. Cold Temperate Grassland 142, Plains Grassland 142.1, Mixed "Short-Grass" Series 142.13 was closest thing--and that missed it--in Brown et al. (1998, p. 40). No appropriate SRM unit for this badlands mixed prairie: the badlands grassland mosaic described (albeit briefly) in Barker and Whitman (1989) was not included in Society for Range Management cover types (Shiflet, 1994). Mosaic of Thin Upland (degraded) range site and Shallow Clay range site (Soil Conservation Service, 1987). White River Badlands Ecoregion- 43h (Bryce et al., 1996).

69. Ultimate on the uplands-Northern Mixed Prairie on interior uplands (Thin Upland was the predominant range site) of the White River Badlands. These two views presented the epitome of grasslands in the Northern Great Plains. Several soil series were represented in the landscape-scale scene provided by the first slide. The land seen in the first photograph was "healed badlands", formerly eroded ravines and canyons that had "haired over" with a cover of green needlegrass, needle-and-thread, sideoats grama, blue grama, and red threeawn (Aristida longiseta) as the major grasses. There was some little bluestem, but this midgrass was uncommon. There were no plants of tallgrasses such as prairie sandreed or big bluestem. This was a mixed prairie of mid- and shortgrasses. Plains yucca (Yucca glauca) was the most abundant forb with silverleaf scurf-pea (Psoralea argophylla) the associate forb species.

Predominant soil on the landscape shown in the first slide was Norka silt loam, a deep soil high in fine sand content and of neutral to moderate alkalinity.

Grassland vegetation presented in the second slide included green needlegrass, needle-and-thread, red threeawn, blue grama, buffalograss, and sideoats grama as the major grasses. In contrast to the range plant community presented in the first slide grassland vegetation in this second slide included some western wheatgrass and lesser amounts of prairie sandreed. Thus it included a more mesic midgrass and some individuals of a tallgrass species. This latter did not, however, exist with enough cover to constitute a tallgrass layer. As with the grassland plant community in the first photograph the range community in the second slide was mixed prairie made up only of mid- and shortgrasses. Plains yucca and silverleaf scurfpea were the dominant and associate forb species, respectively.

Soils of grassland shown in the second slide included Fairburn, a loam to a gravelly clay loam that generally consist of shallow soils formed on steep slopes and that are 18 to 35% clay with moderate to strong alkalinity and Midway which are shallow soils derived from platy shale and that are 35 to 45% clay being moderate to strongly alkaline.

These two views provided textbook examples of Northern Mixed Prairie.An interesting and diagnostic thing visible in both of these slides was presence of capsules remaining on yucca plants. The blooms of Yucca species are so palatable to grazers like cattle and buffalo that presence of so many capsules was a foolproof sign that there had been little to only light grazing of this grassland by bovids.

Badlands National Park, Pennington County, South Dakota. July. FRES No. 38 (Plains Grasslands Ecosystem). K-61 (Wheatgrass-Grama-Buffalograss) was generic at type (versus range site) scale for White River Badlands. Cold Temperate Grassland 142, Plains Grassland 142.1, Mixed "Short-Grass" Series 142.13 was closest thing--and that missed it--in Brown et al. (1998, p. 40). No appropriate SRM unit for this badlands mixed prairie: the badlands grassland mosaic described (albeit briefly) in Barker and Whitman (1989) was not included in Society for Range Management cover types (Shiflet, 1994). Thin Upland range site (Soil Conservation Service, 1987). White River Badlands Ecoregion- 43h (Bryce et al., 1996).

Going, going, gone- The next three slides showed an erosional sequence using three contemporary (current) stages of weathering of spires, pinnacles, or towers (the original shape and size of any of the three badlands structures was obviously unknown) within the White River Badlands. These three location were on the upper or northern grassland of the White River Badlands (north of the Badlands Wall). Surrounding each of the three, current, weathered earth structures was microtopography none of which was identical to any one or both of the other two. Therefore, each of the three grassland communities was on a different range site or, more precisely, a different mosaic of range sites. The range sites or, again, the pattern or complex of range sites were results of different stages of erosion. They were, to some degree, stages in land formation in conjuction with differences in soils or stages in soil formation (genesis). Development of grassland vegetation progressed parallel to that of soil and land formation. All three of these grassland plant communities were interpreted as climax or potential natural vegetation for their respective range sites or, more correctly, mosaic or complex of range sites.

70. Wearing down- Remains of a portion of a previous tower, spire, pinnacle within White River Badlands This and the next two subsequent slides provided a classic example of the interrelations of geologic erosion, soil genesis and plant succession on Northern Mixed Prairie. The major range plants on this swatch of badlands werer western wheatgarss, needle-and-thread, green needlegrass (in that rough order of cover and phytomass) with blue grama being a distant fourth yet with enough cover density, etc. to compirse a sporadic shortgrass layer. Clearly this rangeland plant community was predominantly one of cool=season (and festucoid) grasses. No tallgrass species were present in this grassland vegetation.

Badlands National Park, Pennington County, South Dakota. July. FRES No. 38 (Plains Grasslands Ecosystem). K-61 (Wheatgrass-Grama-Buffalograss) was generic at type (versus range site) scale for White River Badlands. Cold Temperate Grassland 142, Plains Grassland 142.1, Mixed "Short-Grass" Series 142.13 was closest thing--and that missed it--in Brown et al. (1998, p. 40). This local range plant community was SRM 607 (Wheatgrass-Nweedlegrass) within the overall badlands grassland mosaic that was described briefly in Barker and Whitman (1989) and not included in Society for Range Management cover types (Shiflet, 1994). Silty range site (foreground) and Badlands Overflow range site (background) (Soil Conservation Service, 1987). White River Badlands Ecoregion- 43h (Bryce et al., 1996).

71. Wore down some more- Textbook example of mixed prairie at base of a former pinnacle, tower, spire, etc. in White River Badlands that had been eroded down to its still visible base. A prominent depression had formed at this remnant of what was probably a majestic badlands earthern form. This recessed surface of the land created a mesic microtopograpy on which a mixed prairie had developed that included tallgrass species (prairie sandreed and little bluestem), misdgrass species (western wheatgrass, needle-and-thread, and green needlegrass), and shortgrass species (blue grama and buffalograss). The three midgrasses were the overall tri-dominants. As was the case for the grassland vegetation in the immediately preceding photograph this range vegetation was composed principally of cool-season (and festucoid) grasses.

Badlands National Park, Pennington County, South Dakota. July. FRES No. 38 (Plains Grasslands Ecosystem). K-61 (Wheatgrass-Grama-Buffalograss) was generic at type (versus range site) scale for White River Badlands. Cold Temperate Grassland 142, Plains Grassland 142.1, Mixed "Short-Grass" Series 142.13 was closest thing--and that missed it--in Brown et al. (1998, p. 40). No appropriate SRM unit for this badlands mixed prairie: the badlands grassland mosaic described (albeit briefly) in Barker and Whitman (1989) was not included in Society for Range Management cover types (Shiflet, 1994). Silty range site (foreground) and Badlands Overflow range site (background) (Soil Conservation Service, 1987). White River Badlands Ecoregion- 43h (Bryce et al., 1996).

72. Wore plum down or a pinnacle no more- A former pinnacle or tower within the White River Badlands that was about as eroded as it could be and still shown some remains or residue of its former form. Major range plants species around this remaining bare patch of a pinnacle were western wheatgrass, blue grama, and buffalograss. Was this the local, range site-specific, climax vegetation or would further geologic erosion of what little of this pinnacle was left progress to another range site with another potential natural (= climax) vegetation? The well-read student of succession could certainly (and clearly) see the polyclimax theory (Tansley, 1935) and the climax pattern theory (Whittaker, 1953) by viewing the local plant communities, the "patches and pieces" of the vegetational mosaic, of the White River Badlands as smaller-scale climaxes based on soil series, slope, aspect, etc. The well-versed and open-minded student could just as easily see the monoclimax theory (Clements, 1916, 1935) by viewing the many local plant communities as merely long-term seral stages corresponding to the various stages of geologic erosion until the White River Badlands are eroded down to a hypothetical peneplain. The successional view depends strictly on the time frame--the temporal scale--at which the dynamic vegetation of the dynamic badlands landscape is interpreted. Or, same thing said another way, one's view of plant succession on the White River Badlands depends on whether the student of vegetation regards 1) spatial scale primary and temporal scale secondary (polyclimax and climax pattern theories) or, alternatively, 2) temporal scale primary and spatial scale secondary (monoclimax).

This author heard the voices of all three renowned successional theorists espousing his unwavering interpretation across these grasslands. Or maybe it was just the wind moving through the ever-eroding badlands.

Badlands National Park, Pennington County, South Dakota. July. FRES No. 38 (Plains Grasslands Ecosystem). K-61 (Wheatgrass-Grama-Buffalograss) was generic at type (versus range site) scale for White River Badlands. Cold Temperate Grassland 142, Plains Grassland 142.1, Mixed "Short-Grass" Series 142.13 was closest thing--and that missed it--in Brown et al. (1998, p. 40). This local range plant community was SRM 609 (Wheatgrass-Grama) within the overall badlands grassland mosaic that was described briefly in Barker and Whitman (1989) and not included in Society for Range Management cover types (Shiflet, 1994). Shallow Clay range site (Soil Conservation Service, 1987). White River Badlands Ecoregion- 43h (Bryce et al., 1996).

73. Mappers' nightmare- Cattle range on mixed prairie with a patchwork of range sites in the interior of the White River Badlands. Across much of the badlands landscape there are numerous range sites--no to mention microsites--on comparatively small acerages. In the lingo of Landscape Ecology this is habitat heterogenity at fine (small) spatial scale. Differential rates of erosion resulting from different resistances to erosion in parent materials and soils combined with varied land surfaces (differences in steepness, aspect, slope) results in micro-mosaics of range environments. This small-scale heterogenity results in remarkable diversity of grassland environments for plant and animal life.

Range sites within the fields of view of these two slides included Silty, Shallow Clay, and Badlands Overflow: Soil series in these views included Orella, Interior, Cedarpass. This makes for a nightmare (or worse) for those mapping soils and vegetation, but it also makes for a wide array and rich assortment of soil-vegetation unitsalong with species richness and biological diversity. At least this is so by standards and spatial scale of regional or zonal grasslands

Major range plant species in the badlands mixed prairie shown here included a mixture of cool-season grasses (western wheatgrass, green needlegrass, needle-and-thread, Junegrass) and warm-season grasses (blue grama, sideoats grama, buffalograss, inland saltgrass [Distichlis stricta= D. spicta var. stricta], and little bluestem). Little bluestem was more of the height and size of a midgrass being more like sideoats grama and western wheatgrass than big bluestem, switchgrass, and prairie sandreed which, while common on some badlands range sites, were absent from this grassland community. This was a mixed prairie comprised of mid- and shortgrass species without a major tallgrass species or major forb or shrub species (ie. no component of these groups of range plants).

Nice herd of Angus or Angus-cross cows and their heavy calves on this allotment of public range.

Buffalo Gap National Grasslands, Pennington County, South Dakota. July. FRES No. 38 (Plains Grasslands Ecosystem). K-61 (Wheatgrass-Grama-Buffalograss) was generic at type (versus range site) scale for White River Badlands. Cold Temperate Grassland 142, Plains Grassland 142.1, Mixed "Short-Grass" Series 142.13 was closest thing--and that missed it--in Brown et al. (1998, p. 40). No appropriate SRM unit for this badlands mixed prairie: the badlands grassland mosaic described (albeit briefly) in Barker and Whitman (1989) was not included in Society for Range Management cover types (Shiflet, 1994). Mosaic of range sites: Silty , Shallow Clay, and Badlands Overflow (Soil Conservation Service, 1987). White River Badlands Ecoregion- 43h (Bryce et al., 1996).

74. Sod table, salt, and saltbush- Example of the sod table micro-land form with saline soil and Nuttall's or Gardner's saltbush (Atriplex nuttallii= A. gardneri= A. tridentata). Sod tables are small up to rather large areas of grass-covered land remaining after erosion removed surrounding portions of land. Sod tables are the relicts or 'all that is left" of a recent (in geologic time) higher land surface. Tough, fibrous roots of grass or, in a few cases, of sedge (eg. threadleaf sedge) effectively held grassland soil against the geologic agents of erosion while unprotected (less protected, anyway) conterminous (contiguous) land was washed, blown, or rolled away by gravity. Sod tables are pedestaled land at larger micro-or meso-scale. Did man's use (maybe abuse or misuse) of the land accelerate soil loss (soil erosion) or was this geologic (natural) erosion? Obviously, some soil series and steeper land surfaces are more prone to soil erosion.

The soil within this Badlands mapping unit appeared to be of the Interior series. This soil mapping unit was described as consisting " of deep, well-drained soils formed in sodium-rich loamy, silty alluvium on fans and level plains" (Soil Conservation Service, 1987, p. 89). Barker and Whitman (1989) explained that in Northern Great Plains badlands accumulations of sodium are widespread, especially on eroded slopes, footslopes and benches at base of slopes.

The range plants in this photograph were yellow wild buckwheat (Ergonium flavum), low shrubs in front or foreground, and Nuttall's saltbush, at right in midground. Nuttall's saltbush was covered immediately succeeding) slide-caption set. Yellow wild buckwheat was at post-bloom stage.

Buffalo Gap National Grasslands, Pennington County, South Dakota. July. FRES No. 38 (Plains Grasslands Ecosystem). K-61 (Wheatgrass-Grama-Buffalograss) was generic at type (versus range site) scale for White River Badlands. Cold Temperate Grassland 142, Plains Grassland 142.1, Mixed "Short-Grass" Series 142.13 was closest thing--and that missed it--in Brown et al. (1998, p. 40). No appropriate SRM unit for this badlands mixed prairie: the badlands grassland mosaic described (albeit briefly) in Barker and Whitman (1989) was not included in Society for Range Management cover types (Shiflet, 1994). Badlands Overflow range site (Soil Conservation Service, 1987). White River Badlands Ecoregion- 43h (Bryce et al., 1996).

75. Salt and saltbush by the table-There are several Atriplex species in the badlands including annual herbs to shrubs. Nuttall's--or is it Garener's saltbush?--saltbush was described by the Great Plains Flora Association (1986, p. 164) as a "highly variable species". Perhaps for this reason there remains considerable confusion regarding proper name of this woody Atriplex species. There are only four woody Atriplex species in the Great Plains Region (Great Plains Flora Association (1986, p. 164). The only other one of these in the White River Badlands was fourwing saltbush (A. canescens) which the species presented here most certainly is not.

Welsh et al. (1987, p. 132) explained that A. gardneri has priority over A. nuttallii (hence it would be Gardner's saltbush) for a species found in the Great Basin as well as on saline habitats of the western Great Plains. They described A. gardneri as "a widely distributed complex of integrading genotypes of great plasticity" with diploid and polyploid (up to at least hexaploid or greater) forms and six taxonomic varieties in Utah (Welsh et al., 1987, p. 132).

Many of the major range plants on the saline, calcareous soils of the White River Badlands are the same species that are dominant, widespread, or common in the Great Basin, including the Great Basin Desert. These include shadscale (Atriplex confertifolia), Nuttall's saltbush, black greasewood (Sarcobatus vermiculatus) , winterfat (Eurotia lanata), big sagebrush (Artemisia tridentata subsp.tridentata), and seablite (Suaeda fruticosa). Inland saltgrass is often the dominant herbaceous species on such alkaline, saline soils. (Recall this grass from above)

Buffalo Gap National Grasslands, Pennington County, South Dakota. July

76. Gardner's or Nuttall's saltbush- Leaders and immature female flowers of A. gardneri (=A. nuttallii= A. tridentata) on the plant that was introduced in the two preceding slides. This is a dioecious species with this being a salty lady that was about to burst into flower.

Buffalo Gap National Grasslands, Pennington County, South Dakota. July

77. Breaks grassland- Landscape of wheatgrass-needlegrass mixed prairie at upper elevation and outskirts of the Yellowstone River Breaks in the unglaciated Northern Great Plains. Western wheatgrass and needle-and-thread were climax co-dominants with Wyoming big sgebrush (Artemisia tridentata subsp. wyomingensis) the major woody species on this range of Northern Mixed Prairie. On some portions of this livestock-excluded range silver sagebrush (A. cana) was the most abundant shrub species. Other important grasses included buffalograss and, with lesser cover locally, blue grama. There was some Kentucky bluegrass, Japanese chess or Japanese brome, cheatgrass or downy brome (much less of this species than Japanese brome) and Sandberg's bluegrass in locally disturbed microsites such as along the pasture road. Grasslike plants were represented by threadleaf caric sedge (Carex filifolia), a local associate and important forage species. Shrub species in addition to Wyoming big sagebrush and silver sagebrush included plains pricklypear, soapweed yucca (Yucca glauca), and the suffrutescent (subshrub), fringed sagebrush (Artemisia frigida). Forbs included Gunnison's sego or mariposa lily (Calochortus gunnisonii), scarlet globemallow, and the naturalized Eurasiaon goatsbeard or salsify (Tragapogon dubia).

Western wheatgrass and needle-and-thread occurred as both 1) consociations of nearly single species composition and 2) mixed co-dominant stands. Wyoming big sagebrush was found as both individual plants and as local, small patches. In this respect, there were areas of this range that at local scale could be interpreted as being of the rangeland cover type, western wheatgrass (SRM 61). This author regarded such a restricted view as a strained interpretation, especially given that there were low overflow (swale) range sites exclusively populated by western wheatgrass immediately adjacent to (contiguous with) the wheatgrass--needle-and-thread community on this hill slope habitat (see below).

Similarly, on this range Wyoming big sagebrush was locally abundant and presented an appearance as a local aspect domiinant, but this and the other shrub speceies combined did not constitute enough of a consistent shrub layer for this range vegetation to be interpreted as a savnna.

Long-term livestock exclosure, Fort Keogh Livestock and Range Research Station, Custer County, Montana. Mid-June, late vernal aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-59 (Wheatgrass-Needlegrass). Variant of 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- Missouri River Breaks Ecoregion 43c (Woods et al., 2002).

78. Needlesome co-dominant- Consociation of needle-and-thread near the top and (mostly) on south slope of a low hill in the margin of Yellowstone River Breaks. On this hill habitat there some local patchlike consociations of western wheatgrass as well as areas with both western wheatgrass and needle-and-thread as co-dominants. Below this there was a swale range site that almost was almost exclusively comprised of western wheatgrass (more of an immense population of this one species). Part of that swale range site was visible in the mid and right background of the first slide which provided a landscape-scale perspective.

The second of these two slides featured the upper sward of a needle-and-thread consociation. A plant of goatsbeard or salsify, soapweed yucca, and fringed sage were also included in this second slide that served as more of a "photoquadrant" of this range vegetation.

Long-term livestock exclosure, Fort Keogh Livestock and Range Research Station, Custer County, Montana. Mid-June, late vernal aspect. Needle-and-thread at soft dough phenological stage. FRES No. 38 (Plains Grasslands Ecosystem). K-59 (Wheatgrass-Needlegrass). Variant of 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- Missouri River Breaks Ecoregion 43c (Woods et al., 2002).

79. At its zenith- Northern Mixed Prairie in Yellowstone River Breaks at peak standing crop. This was a consociation of needle-and-thread which was co-dominant with western wheatgrass in this pristine, primarily cool-season grassland. Wyoming big sagebrush, the major shrub species in this range plant community, was present as both single plants and small groups of several plants. Contrary to the aspect afforded in these two photographs, Wyoming big sagebrush was not abundant enough or afford enough foliar (aerial) cover to interpret this range vegetation as savanna. Other shrubs present included plains pricklypear, soapweed yucca, and fringed sage. There was a vegetationally significant shortgrass component comprised of buffalograss and blue grama with the former more abundant (greater cover) than the latter. There was more cover of treadleaf sedge, the only grasslike plant species, than of the two shortgrasses combined, at least at this point in the annual cycle of plant and vegetation growth (= in the vernal society). Forbs included Gunnison's sego or mariposa lily, the most conspicuous, scarlet globemallow, and goatsbeard or salsify (naturalized Eurasian composite). In these two views of this pristine grassland there were no Eurasian grasses present.

These two slides provided nested (telescoped) views of the same grassland location with the second image being an obvious sublocation within the vegetation presented in the first photograph. These two images and all except two other slides of this particular livestock-excluded-from range where taken under full-sunlight conditions. The two exceptions were the two photographs in the next slide-caption set.

80. At its zenith under an overcast sky- Northern Mixed Prairie in Yellowstone River Breaks at peak standing crop. Pair of nested (telescoped) "photoplots" of a needle-and-thread--western wheatgrass (co-dominants) mixed prairie with Wyoming big sagebrush as the major woody species. This pair of images duplicated the two in the immediately preceding set, except that these two photographs were taken under an overcast sky when a cumulus cloud blew between the sun and photographic subject. The first of these two slides was a nearly exact copy of the first image in the preceding slide-caption set (ie. taken from almost the identical camera location).

The purpose of this experimental exercise was to present to viewers the differences--subtle though they were--between a full-sun and overcast shot. Viewers will note that contrary to oft-heard claims about superiority of slightly overcast shots, the bright, no-shade photographs were generally more revealing of details ranging from clairty and definition of cumulus clouds to presence of Gunnison's mariposa or sego lily in the grassland sward. Students should also note that there was greater contrast between colors of grass and sagebrush in sunlite shots. However, panicles and tillers of needle-and-thread were more discretely visible (discernible) in the overcast shots.

Consistent with improved capability of the human eye to discern grass shoots in overcast shots was distinction between needle-and-thread and western wheatgrass in the second (zoomed-in) shot in this set. This was not overwhelmingly or completely the case however as this second photograph was taken at closer camera distance to the vegetation. Comparison of this second slide with the second slide in the preceding set proved the fact of closer camera focal point which obviously confounded this little photographic experiment.

In the second of these slides nonflowering grass shoots from the lowermost left corner extending diagonically to Wyoming big sagebrush were of western wheatgrass. Petals of Gunnison's mariposa lily were present in immdeiate center foreground of this same photograph.

81. Array of phytocharacters- Group of major plant species of a needle-and-thread--western wheatgrass dominated Northern Mixed Prairie on the south slope of a low hill in the Yellowstone River Breaks. This "photoplot" was taken to include a small (local) group of Wyoming big sagebrush plants so as to include the major woody species of climax range vegetation that was--in spite of this featured sagebrush group--clearly grassland and not grass-shrub savanna. Wyoming big sagebrush was, nonetheless, an associate species. Needle-and-thread was at peak standing crop in soft-dough grain stage while western wheatgrass was prebloom (mid-boot) stage of phenology. Threadleaf caric sedge was present at such cover and biomass as to be an associate herbaceous species (shown in slides below). The other shrub in this "group shot" was plains pricklypear. There were two plants of Gunnison's mariposa or sego lily, one of the major forbs on this range.

Structure and composition of this climax range plant community were presented in this composite "photoquadrant".

This photograph was taken under full sunlight in contrast to two views of this climax vegetation taken under an overcast sky (cumulus clouds between sun and range plant community) and presented in the immediately preceding two slides. As a follow-up to above comparisons of full-sun versus overcast photographs, viewers should note that needle-and-thread and western wheatgrass can be readily distinguished in this photograph taken under intensely bright light. Viewers should also note that, contrary to frequent statements,there were no shadows present in this full-light photograph. Same held true for subsequent slides taken under intense light of full sun. Next slides please...

82. Phytoactors on a Northern Great Plains range stage- Major range plant species on a grassland of Northern Mixed Prairie that developed on a south slope in the Yellowstone River Breaks. These two "photosamples" of this climax vegetation contained included plants of the following species (in one or both slides): needle-and-thread (peak standing crop; soft-dough grain phenological stage), western wheatgrass (mid-boot phenological stage), threadleaf sedge (peak standing crop; mature achene stage of phenology), plains pricklypear (immediate prebloom stage), Gunnison's mariposa or sego lily (full-bloom phenological stage), and scarlet globemallow (nearly adult size; various prebloom phenological stages).

83. More phytoactors on this range stage- Portion of climax mixed prairie vegetation on which needle-and-thread and western wheatgrass were co-dominants. Seen here was a needle-and-thread consociation that developed on the south slope of a low hill in the Yellowstone River Breaks. Threadleaf caric sedge was the associate species in photographic sample of the range plant community.

This photoquadrant" featured one plant of Wyoming big sagebrush (large shrub on left) and fringed sage or sagebrush (small shrub on right). Silver sagebrush was a third Artemisia species growing in this climax vegetation. There was not a plant of silver sagebrush growing close enough to make it into this "staff photograph".

This pristine example of climax mixed prairie had developed in the unglaciated portion of the Northern Great Plains.

84. Spiny beauty on the northern plains- Plains pricklypear (Opuntia polycantha) on Northern Mixed Prairie co-dominated by needle-and-thread and western wheatgrass. Needle-and-thread was the local hilltop dominant on this margin of the Yellowstone River Breaks of the sedimentary Northern Great Plains.

Long-term livestock exclosure, Fort Keogh Livestock and Range Research Station, Custer County, Montana. Mid-June; immediate pre-bloom to full-bloom phenological stages of plains pricklypear.

85. Bird's eye (of a low-flying bird) view- Top-down visual perspective of the sward of Northern Mixed Prairie on drier south slope in Yellowstone River Breaks. Major species were western wheatgrass, needle-and-thread, and threadleaf caric sedge, and (in that order of relative foilar cover). Students should locate some of each of these species in this "photoplot". Also present (though more conspicuous than being a meaningful component to cover, let alone biomass) was a plant of Gunnison's sego or mariposa lily in peak bloom.

(The black spot below the flower of the mariposa lily is its shadow. In keeping with the full-sun versus overcast photographs discussion above it was shown that in this and other slides that with narrow shoots of graminoids and smaller shrubs there is tyically minimal or no shadows at all in photographs taken with bright, full sun directly overhead. Hence, bright, sunlite photographs work well for most grassland vegetation.)

86. One to make Solomon envious- Gunnisons's mariposa lily or sego lily (Calochortus gunnisoni) growing on pristine Northern Mixed Prairie in the unglaciated Nolrthern Great Plains with needle-and-thread, western wheatgrass, threadleaf caric sedge, buffalograss and blue grama as the manjor garminoids along with the shrubs Wyoming big sagebrush, plains pricklypear, soapweed yucca, and fringed sage.

No sane range observer could question the sheer beauty and distinctiveness of this range forb. Gunnison's sego lily is one of the more widely distributed Calochortus species with a species range from British Columbia and Alberta to Nebraska and south to Arizona. On this Excellent condition class mixed prairie prairie Gunnison's mariposa lily was never abundant as it grew as isolated individual plants or, at best, groups of two or three plants. Color of this species varies from all-white to petals with pronounced yellowish purplish coloration (Hitchcock and Cronquist, 1973, p. 686). Definitive source on Calochortus species was that of Gerritsen and Parsons (2007; see especially ps. 160-161 for C. gunnisoni). This labor-of-love scholarly acheivement is a beautiful book; highly recommended.

Long-term livestock exclosure, Fort Keogh Livestock and Range Research Station, Custer County, Montana. Mid-June, peak bloom stage.

87. Cow's eye (of a Line One Hereford) view- Two views--at approximate level of a cow moving from grazing point to another--of the sward of Northern Mixed Prairie on the south slope of a low hill in the margin of the Yellowstone River Breaks. Major species were needle-and-thread-, buffalograss, threadleaf sedge, and western wheatgrass (in that order of relative foilar cover). Students should practice their plant identification skills by finding some of each of these range species in this "photosample". There was some foliar cover of blue grama in the second of these photographs, but only in minor quantities in this late vernal vegetational society.

88. Grassland ecotone- Transition zone between an upland site of mixed prairie co-dominated by needle-and-thread and western wheatgrass (foreground of first slide, background of second slide) and a swale site dominated (almost exclusively) by western wheatgrass (background of first slide, foreground of second slide). By way of explanation, directions of camera orientation were reversed in these two photographs (hillside facing downslope versus base of hillslope facing uphill in first and second slides, respectively). Needle-and-thread rated somewhat superior to western wheatgrass in the co-dominated mixed prairie (which had a meaningful shortgrass component) whereas western wheatgrass was the sole dominant on the swale or overflow habitat (with buffalograss and blue grama being associate, though marginally so, species).

Climax vegetation of the needle-and-thread--western wheatgrass community was detailed in the last (immediately preceding) eight slide-caption sets going from landscape-scale down to local or microsite-scale of the sward.

Long-term livestock exclosure, Fort Keogh Livestock and Range Research Station, Custer County, Montana. Mid-June, late vernal aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-59 (Wheatgrass-Needlegrass). Ecotone of 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- Missouri River Breaks Ecoregion 43c (Woods et al., 2002).

89. Patch in an ecotone- Local (small spatial scale) consociation of western wheatgrass within a transition zone between mixed prairie co-dominated by needle-and-thread and western wheatgrass and a swale (overflow) range site dominated solely by western wheatgrass. Even within the comparitively narrow ecotone between these two rangeland cover types (SRM 607 and SRM 610) there small patches populated almost exclusively by one or the other of these festucoid grasses. This was more commonly the case for the rhizomatous (sod-forming) western wheatgrass than the cespitose (tufted or bunchgrass) needle-and-thread.

Within this single-species dominated sward there were scattered plants of Wyoming big sagebrush and silver sagebrush. There was also a comparatively large plant of winterfat (Eurotia lanata) in the lefdt margin.

Long-term livestock exclosure, Fort Keogh Livestock and Range Research Station, Custer County, Montana. Mid-June, late vernal aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-59 (Wheatgrass-Needlegrass). Ecotone of 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- Missouri River Breaks Ecoregion 43c (Woods et al., 2002).

90. Co-dominants in an ecotone- Needle-and-thread and western wheatgrass in a narrow ecotone between an upland mixed prairie co-dominated by needle-and-thread and western wheatgrass and a lowland (= flat or swale) dominated exclusively by western wheatgrass. In these two "photoplots" the strongly rhizomatous western wheatgrass was the dominant and the strictly cespitose needle-and-thread was the associate species.

The co-dominated mixed prairie with threadleaf sedge and buffalograss with locally numerous plants of Wyoming big sagebrush and silver sagebrush was covered above. The western wheatgrass flat was covered the two slide-caption set immediately below.

Long-term livestock exclosure, Fort Keogh Livestock and Range Research Station, Custer County, Montana. Mid-June, late vernal aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-59 (Wheatgrass-Needlegrass). Ecotone of 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- Missouri River Breaks Ecoregion 43c (Woods et al., 2002).

91. One major species of grass (with one major grazer)- Landscape-scale views from upper (slightly higher elevation) to lower limits of a swale range site populated almost exclusively by western wheatgrass and with black-tailed prairie dog as the overwhelmingly dominant herbivore consumer. Western wheatgrass, by the simple nature of this grassland essentially being a a single-species consociation, was the keystone plant species (producer). This was a cattle range that was also grazed by deer, but at time of these photographs prairie dogs were clearly the keystone animal species (consumer). In fact, herbivory by black-tailed prairie dogs had driven management of this grazing unit for the last several years.

There was a story behind the story to range vegetation seen here. Off the record, range technicians responsible for this experimental pasture on the Fort Keogh experimental range explained to the author that this prairie dog town (= population or colony) began when a research leader used this pasture as a heavy stocking rate treatment for one of his grazing trials. Some prairie dog control had been conducted on this western wheatgrass-dominated grassland a few years prior to time of these slides. This was yet another example of the general finding that black-tailed prairie dogs are often the result and not the source (effect not cause) of a heavily grazed range and any impacts resulting therefrom. It was also revealing--though of what may be known but to God--that at this point in life (time span) of the prairie dog town (and with some human control of prairie dog numbers) the keystone consumer species had not been influenced botanical composition of the range plant community, at least not in any substantitative way given that vegetation was still a western wheatgrass consociation.

Clarification of climax vegetation classification: strictly speaking this single midgrass species-dominated range plant community could be interpreted as either true prairie (Clements,1920, ps. 122-124) or mixed prairie (Clements, 1920, pgs. 137-138) as it is a dominant species fnd forms consociations in both major grassland associations.

Long-term livestock exclosure, Fort Keogh Livestock and Range Research Station, Custer County, Montana. Mid-June, late vernal aspect. FRES No. 38 (Plains Grasslands Ecosystem). K-59 (Wheatgrass-Needlegrass). SRM 610 (Wheatgrass-Needlegrass). Cold Temperate Grassland142- Plains Grassland 142.1, but no appropriate Series in Brown et al. (1998, p. 40). Northwestern Great Plains- Missouri River Breaks Ecoregion 43c (Woods et al., 2002).

Northern Mixed Prairie-Palouse Prairie Ecotone or Transition Grassland

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

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

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

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

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

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

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

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

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

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

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

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

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

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

96. Wide and widespread silver- Two plants of silver (or silvery) or tailcup lupine (Lupinus argenteus var. argenteus) on Palouse Prairie-Northern Great Plains transition grassland in central Montana. Silver lupine is one of the most widely distributed Lupinus species in North America extending from the Southern Great Plains of the Plains States up through the Prairie Provinces and British Columbia. Silver lupine is adapted to a wide array of habitats and grows on grasslands, forests, parklands, sagebrush-grass savannahs, and up to alpine (Hermann, 1966, ps. 103-104; Lesica, 2012, p. 310). In keeping with this wide adaptation, a number varieties and great variability within the species (Great Plains Flora Association, 1986, p. 463) have been reported for L. argenteus. Lesica (2012, ps. 310-311) described four varieties in Montana. Some authorities (eg. Burrows and tyrl, 2013, ps. 571-572) include L. caudatus in the L. argenteus group.

Silvery lupine has a large caudex or proaxis (a semi-woody base of herbaceous plants; the union of root and shoot near or slightly below the soil surface) from which a number of shoots arise (Great Plains Flora Association, 1986, p. 463; Lesica, 2012, p. 310). This morphological festure was conspicuous in the two above photographs.

The Lupinus genus is a challenging one even (maybe, especially) for taxonomists given integradation and variation within the large number of species. Lupinus is also one of the more important group of species from standpoint of sheep poisoning on the range. Although Lupinus species furnish forage for livestock and wildlife (Hermann, 1966, 103) a number of species of this papilionaceous legume can be highly toxic. Borrows and Tyrl (2013, ps. 567- 576) provided the most recent definitive treatment for Lupinus toxicity (at time of this writing) and noted that there are approximately 150 Lupinus species (not to even begin with subspecies and varieties) in North America most of which have not been been studied with ragard to poisonous principles.

The toxic principle in North American Lupinus species is a group of quinolizidine alkaloids, the main two of which are lupanine and anagyrine, that cause acute neurological, hepatic, and renal disorders--a classic syndrome--and, in some instances, a reproductive malady known as crooked calf disease that induces abortion, or deformed fetuses. The disease-causing substances, the alkaloids, are produced through chemical transformation by a fungus (Phomopsis leptostromiformis= Diaporthe toxica). Lupinosis, the generic term for Lupine-induced toxicity, is most serious in sheep, but it also affects goats, cattle, and horses. Tetragenic effects are most important in cattle. order Again, Borrows and Tyrl (2013, ps. 567- 576) is the definitive source though, of course, Kingsbury (1964. ps. 333-341), while limited by its earlier date, will always be an old, revered standby. In fact, the quinolizidine alkaloids had been implicated as the causative principle even back at the time of Kingsbury (1964, p. 333-334).

L. argenteus contains both alkaloid types--lupanine and anagyrine--and is one of the more widely studied, including feeding trials, poisonous lupines (Burrows and tyrl, 2013, ps. 571-575) in regards to livestock poisoning. Strickingly beautiful, but deadly under certain conditions. One of the main one of these is sheep consuming mature or maturing legumes (fruits) of lupines or when hungry and being driven through lupine-endowed ranges.

Meagher County, Montana. Late June; full-bloom phenological stage.

97. Silvery shoots- Shoots, including leaves and infloresecences, of silvery lupine growing on lush bluebunch wheatgrass-Idaho fescue transition from Palouse Prairie to Northern Great Plains mixed prairie--grassland in the dissected plains area of northcentral Wyoming. Lesica (2013, p. 310) interpreted the inflorescence of Lupinus species as a raceme.

Sheridan County, Wyoming. Mid-June; peak-bloom phenological stage.

98. Silverish green- This sunlite view of the spreading leaves with their six to eight leaflets of silvery lupine showed off the basis of the common name for this species. This was one of several plants growing on a bluebunch wheatgrass-Idaho fescue-dominated grassland in the dissected plains portion of the Northern Great Plains in northcentral Wyoming.

Sheridan County, Wyoming. Mid-June.

99. Flowers and fruits in the hills- Papilionaceous flowers along a portion of a raceme (upper slide) and just-starting to -ripen legumes (second slide) of silvery lupine growing with bluebunch wheatgrass and Idaho fescue of a transition--from Palouse Prairie to Northern Great Plains mixed prairie--in the dissected plains area of the Northern Great Plains in northcentral Wyoming.

The legumes, especially their seeds, are one of the most poisonous parts of this beautiful and, too often, deadly range forb.

Sheridan County, Wyoming. Mid-June.

Northern Mixed Prairie Savanna or Northern Mixed Grass-Shrub Steppe

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

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

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

100. 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).

101. 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.

102. 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.

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

Interior Pacific Northwest Mixed Prairie

108. Northern Great Plains mixed prairie-Aligned coulees along Old Man River (due to Chinook winds causing greater snowmelt on south slopes so less mesic vegetation and relatively greater geologic erosion) in southern Alberta.Wheatgrasses (western wheatgrass and slender wheatgrass [Agropyron trachycalum] are dominant), green needlegrass, blue grama, Indian ricegrass, Junegrass, basin wildrye (Elymus cinerus), and Sandberg bluegrass (Poa sandbergii = P. secunda) plus Astragalus spp. Prominent shrubs are skunkbrush sumac and chokeburry (Prunus virginiana).

The author interpreted this grass as mixed prairie given a prominent tallgrass component in basin wildrye and, arbitrarily, green needlegrass along with a remarkably diverse species composition. Rationally this grassland vegetation could about as easily have been interpreted as true prairie (see clarifying ecological and organizational note shortly below.

University of Lethbridge campus (the most beautiful I was ever on). July. FRES No. 38 (Plains Grasslands Ecosystem). K- 59 (Wheatgrass-Needlegrass) based on dominant species but similar to K- 57 (Gramagrass-Needlegrass-Wheatgrass). No SRM listing for this widely distributed form of northern mixed prairie.

109. Basin or giant wildrye- This is one of the most important and productive grasses in the Intermountain Region, especially the Great Basin, and Northern Great Plains. Once a major source of hay for U.S. Cavalry horses this large and beautiful bunchgrass was overcut along streams and bottomlands until became (and remains) relatively rare. University of Lethbridge, Alberta. July.

110. Inflorescence (spike) of basin wildrye- University of Lethbridge, Alberta, July.

111. Showy milkweed (Asclepias speciosa)- Showy milkweed is one of the more common representatives of Asclepias found on the mixed prairie of the Northern Great Plains. There are at least 16 species of milkweed in North America that have been documented to cause livestock poisoning. Students are referred to the classic text of Kingsbury (1964) and the recent encyclopedic reference by Burrows and Tyrl (2001, ps.125-135 passim). The inflorescences missing from shoots in the lower left foreground were apparently grazed by mule deer based on tracks in the mud adjacent to the plant.

Cardston Municipal District, Alberta. July.

112. Flower cluster of showy milkweed- Study shot of one inflorescence of Asclepias speciosa. Otero County, Colorado. Late June.

Western Wheatgrass-Dominated Great Plains Grasslands and a Few "Odd Lots" of Mixed Prairie Range Types

Western wheatgrass (Agropyron smithii) is one of the more widely distributed midgrasses (grass species of intermediate height in contrast to tallgras and shortgrass species). Yet many, probably most, of the rangeland cover types dominated by western wheatgrass tend to be more mesic and, with important exceptions such as SRM 610, Wheatgrass, (Shiflet, 1994), tend to be more local or, at least, more restricted in area than mixed prairie range types dominated by other widely distributed grasses such as blue grama, buffalograss, and little bluestem.

Importance of western wheatgrass is perhaps most evident in popular culture and political affairs by its symbolism as the state grass of Wyoming, South Dakota, and North Dakota. Western wheatgrass is a major range plant species throughout much of its biological (species) range which extends across Canada from Ontario to British Columbia and from California to Kentucky and Tennessee and south into Mexico (Fernald, 1950, Correll and Johnston, 1979; McGregor et al, 1986; Barkworth et al., 2004).

With this rationale a separate section of this chapter was devoted to western wheatgrass-dominated range along with some usual range types.

113. Mixed prairie-ponderosa pine (Pinus ponderosa) savanna— Diversegrassland understory of both warm-season grasses (big and little bluestem and upland switchgrass dominant) and cool-season mid-grasses (needle-and-thread [Stipa comata ], green needlegrass [S. viridula ] and Canada or nodding wildrye [Elymus canadensis] dominants).Stoney hills range site.Custer State Park, South Dakota. July. Transition between or mosaic of FRES Nos. 39 (Prairie Ecosystem) and 21 (Ponderosa Pine Ecosystem);K-66 (Bluestem Prairie) and K-16 (Black Hills Pine Forest), respectively. Northern Great Plains variant of SRM 110 (Ponderosa Pine-Grassland). Northwestern Great Plains-Semiarid Pierre Shale Plains Ecoregion, 43g ( Bryce et al., undated).

114. Mixed Prairie of unusual biological diversity: western wheatgrass is the dominant but also needle-and-thread, green needlegrass, Canada wildrye, sideoats grama, blue grama, buffalograss, wild alfalfa.Gallery riparianwoodland of bur oak (Quercus macrocarpa), green ash (Fraxinus pennsylvanica), eastern cottonwood, and a willow (Salix spp.). Clayey range site. Custer State Park, South Dakota. FRES No. 38 (Plains Grasslands Ecosystem) and FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem). K-57 (Gramagrass-Needlegrass-Wheatgrass) with gallery vegetation being K-89 (Northern Flood-Plain Forest). SRM 608 (Wheatgrass-Grama-Needlegrass). Northwestern Great Plains-Semiarid Pierre Shale Plains Ecoregion, 43g ( Bryce et al., undated).

115. Mixed Prairie with a hardwood gallery woodland ecosystem along a creek- In the foreground is a typical consociation of western wheatgrass in a range community dominated by a three-way mix of western wheatgrass, big bluestem, and needle-and-thread. The woodland is bur oak, green ash, eastern cottonwood, and willow with Canada or nodding wildrye as the main understory herb. The grassland is a Clayey range site and the gallery woodland is on an Overflow range site. Floodplain of Lame Johnny Creek, Custer State Park, Custer County, South Dakota. July. FRES No. 38 (Plains Grasslands Ecosystem), K-57 (Gamagrass-Needlegrass-Wheatgrass) and gallery woodland is FRES No. 17 (Elm-Ash-Cottonwood Forest Ecosystem), K-89 (Northern Flood-Plain Forest). SRM 608 (Wheatgrass-Grama-Needlegrass). Northwestern Great Plains-Semiarid Pierre Shale Plains Ecoregion, 43g ( Bryce et al., undated).

116. Consociation of western wheatgrass- This is a natural single-species stand of western wheatgrass on a Dense Clay range site in the Northern Great Plains mixed prairie. There are a few "stray" green needlegrass (Stipa viridula) individuals in this community, but it is a textbook example of the Clementsian consociation (Weaver and Clements, 1938, p. 94 on which western wheatgrass was specified as one of the consociations comprising the true prairie association). Peak standing crop at seed-ripe phenological stage. Estival aspect, July. Pennington County, South Dakota. FRES No. 38 (Plains Grassland Ecosystem), K-57 (Gamagrass-Needlegrass-Wheatgrass), SRM 610 (Wheatgrass). Northwestern Great Plains-Semiarid Pierre Shale Plains Ecoregion, 43g ( Bryce et al., undated).

117. Big sky, big country, and big patch of western wheatgrass- Landscape of Northern Mixed Prairie in the unglaciated (sedimentary) Northern Great Plains of southeastern Montana. Grassland in these two large-spatial scale photographs consisted of at least two range site associations: 1) Clayey and Shallow Clay and 2) Dense Clay-Clayer-Saline Upland, both in 10-14 inch precipittion zone (Ross and Hunter, 1976, ps. 12-14). These added to a sum of four range sites. The predominant range site in foreground of these two images was Dense Clay. Range vegetation on this Dense Clay site consisted almost exclusively of western wheatgrass. There were some local areas of inland saltgrass (Distichlis stricta) and even smaller patches of Sandberg's bluegrass, Kentucky bluegrass, and Japanese chess or Japanese brome. These latter three species usually grew as local populations (single-species stands). Obviously western wheatgrass is a decreaser that is well adapted to heavy grazing, including some overuse, as well as some overgrazing (longterm overuse). The three other grasses were either naturalized Eurasian species so as to be invaders or, in case of Sandberg's bluegrass, a native increaser.

This range was being grazed to a such a high degree of use that there were almost no tillers with spikes (ie. close to zero sexual reproduction on this heavily grazed spring range). There was still some time remaining in the wheatgrass flowering period, but dearth of sexual shoots was due to heavy defoliation proven by abundance of spike-bearing tillers in barrow ditches and fence lines outside this range. Tremendous quantities of grass shoots had been produced in this spring of record snow melt and rainfall. Asexual reproduction of wheatgrass rhizomes and tillers was clearly more than adequate to maintain present populations of western and thickspike wheatgrasses.

It was absence of other species, especially such bunchgrasses as basin wildrye and green needlegrass, that bespoke of past grazing mismanagement, and current state of heavy grazing would definitely not permit persistence--let alone establishment--of these less grazing tolerant, taller-growing decreasers.This degraded range obviously had been abused in the past (ie. had a grazing history that included some mismanagement, including overgrazing). It was currently being grazed at a quite high stocking rate--in fact, probably being overused at present--though this was not conspicuous in the currrent year of record precipitation.

This fundamental lesson in ecology and management of Northern Mixed Prairie range was made even more axiomatic when this and the next overgrazed wheatgrass range were compared to another range of the same range site (Dense Clay) that was in climax state.

Absence of almost all woody plants from this range suggested application of some bursh control practices. For instance, the nonsprouting Wyoming big sagebrush, a common--though often sparsely present--shrub on grasslands adjoining this range is readily killed by 2,4-D. (Shrubs on a similar and adjoining range were shown below).

These two photographs were taken from the same exact camera location within one or two minutes of each other. The first slide was taken under a momentary full-sun condition with a sky full of large cumulus clouds. The second slide was an overcast shot taken when one of these wide-expanse clouds was between the sun and the landscape so as to cast a shadow completely over the rangeland across camera range (28mm lens). The characteristic glaucous-green coloration of western wheatgrass was more visible under the overcast sky. The distinctive bright, light green of inland saltgrass was only detectable and distinguishable from the glaucous coloration of westrn wheatgrass in the full-sun shot. Greater light in the full-sun photograph revealed the ripening (maturing/senescing) sexual tillers of western wheatgrass which were distinguishable from this species' typical glaucous-green coloration still present in leaves. Paired photographs combined to present to viewers a more "balanced perspective" of this mixed prairie vegetation.

Carter County, Montana. Mid-June; late vernal aspect; grain-ripe stage of western wheatgrass. FRES No. 38 (Plains Grassland Ecosystem), K-57 (Gamagrass-Needlegrass-Wheatgrass). Variant of SRM 607 (Wheatgrass-needlegrass). Cold Temperate Grassland 142, Plains Grassland 142.1, but no specific Series unit in Brown et al. (1998, p. 40). Northwestern Great Plains- Montana Central Grasslands Ecoregion 43n (Woods et al., 2002).

118. Mixed prairie range with most species grazed out- Cattle range of Northern Mixed Prairie overgrazed to the extent that grassland vegetation was composed almost exclusively of western wheatgrass in some portions and with thickspike wheatgrass in other portions. Other potential climax (decreaser) grasses of this range plant community that would have been major species had they not been grazed out included green needlegrass and basin wildrye (Elymus cinerus). (An example of this potential, climax, mixed prairie vegetation was presented and described immediately below.) This (and the following example of pristine range) was a Dense Clay range site (Ross and Hunter, 1976, p.13).

This range was currently receiving heavy use up to overuse. There were almost no sexual shoots remaining at time of photographs. Asexual reproduction of these two strongly rhizomatous wheatgrass species appeared to be adequate to maintain current populations (which comprised almost all existing phytomass on this range).

The state of current overuse and, almost assuredly, past overgrazing was shown by abundance of the naturalized, Eurasian annual composite, common dandelion (Taraxacum officinale), and the native, typically weedy composite, western yarrow (Achillea millefolium complex= A. millefolium subsp. lanulosa; traditionally, A. lanulosa).

Continued abundance and domination by western wheatgrass, the climax dominant, "spoke volumes" as to the tolerance of abuse inherent in this midgrass species. Much of this adaptation to heavy grazing (high degree of defoliation) can be explained by its strongly rhizomatous habit. Range site is another factor in persistence of this decreaser under heavy stocking rates. In typical--not to mention wet--springs range sites of slowly permeable soils, such as this Dense Clay range site, have standing water on the land surface from early even into late parts of the western wheatgrass-growing season. Under these conditions ranchers turn out on later on water-indunated range and cattle are less prone to graze such range sites on which animal movement is impeded and made uncomfortable by deep, sticky mud. Yes, such situations do lend protection to associated species such as green needlegrass and basin wildrye, but these cespitose species (bunchgrasses) are less tolerant or resistant (whichever term best applies) than the rhizomatous, sod-forming western wheatgrass. The taprooted, rosette-leafed, achene-rich, ubiquous ruderal composite called dandelion also does quite well under the continued disturbance of overgrazing, thank you very much.

This range was being grazed by large Angus bulls of superior conformation. This was the bull battery pasture for a ranch. High-value bulls like these deserved better of the ranch they lived on. Proper animal husbandry, especially for cattle of superior quality, includes provision of high-quality forage. Outstanding sires notwithstanding, this ranch scene was not an example of admirable beef production.

For an example of proper range management of this same range plant community (plus valuable wetland to boot) please scroll down.

Carter County, Montana. Mid-June; late vernal aspect grain-ripe stage of western wheatgrass. FRES No. 38 (Plains Grassland Ecosystem), K-57 (Gamagrass-Needlegrass-Wheatgrass). Variant of SRM 607 (Wheatgrass-Needlegrass). Cold Temperate Grassland 142, Plains Grassland 142.1, but no specific Series unit in Brown et al. (1998, p. 40). Northwestern Great Plains- Montana Central Grasslands Ecoregion 43n (Woods et al., 2002).

Note on range evaluation: The two overused and, previously, overgrazed ranges described above( to large extent single-species stands of western wheatgrass or of western and thickspike wheatgrass depending on local populations) could be assigned to range condition classes ranging from Fair to Excellent depending on how Soil/Natural Resource Conservation Service guidelines were applied. If all wheatgrass (western and thickspike) as decreaser(s) counted toward climax as permitted in the first National Range Handbook (Soil Conservation Service, 1967) condition class would be Excellent. By methods and standards of the second National Range Handbook (Soil Conservation Service, 1976) arbitrary proportions of plant species could be applied toward toward condition classes of these two ranges could vary from Fair through Good to Excellent. The same situation obtains for the current on-line National Range and Pasture Handbook (Natural Resources Conservation Service, 1997 through present), but by criteria for the state-and-transition model.

From any meaningful application of standards of biodiversity, range type/range site descriptions, on-the-land-experience, and common sense these two mixed prairie ranges converted into grazing monocultures and, in some parts, bicultures of wheatgrass would be far from Excellent condition class though the could perhaps be justified in Good class.

Now for the mint range- the following section was devoted to the climax vegetation of Northern Mixed Prairie on a Dense Clay range site, including an ephemeral wetland. This pristine, herbaceous vegetation on a semiarid, level landscape consisted of the following four communities: 1) western wheatgrass-thickspike wheatgrass-Wyoming big sagebrush savanna, 2) basin wildrye-western wheatgrass grassland, 3) an ecotone between these two plant communities, and 4) wetland herbaceous community dominated by pale spikerush (Eleocharis macrostachya).

This climax range vegetation was an example of one of the more mesic forms or types of mixed prairie on the unglaciated Northern Great Plains. This example served as basis of comparison for the degraded ranges of the Dense Clay range site (Ross and Hunter, 1976, p. 13) presented immediately above.

119. Midgrass-shrub savanna- Wheatgrass-sagebrush savanna on a slowly permeable habitat (Dense Clay range site) in unglaciated (sedimentary ) Northern Great Plains Western wheatgrass was the dominant grass and range species while Wyoming big sagebrush was the associate species on this savanna community within the greater Northern Mixed Prairie. Thickspike wheatgrass was present as the second major grass (associate grass species). Wyoming big sagebrush was the only shrub species that this author could find in this specific range.

This wide-scale view showed viewers the physiogonomy of this climax range savanna. Species composition and structure of this sod(turf)grass-shrub range community were presented in the two immediately succeeding slides. The landscape-scale community (total of four specific climax plant communities) comprised the specific range site (Dense Clay). This wheatgrass-big sagebrush savanna was one of four communities on this particular pasture (grazing unit) which encompassed a fairly large expanse of Northern Great Plains landscape. The other communities (all covered in this section) included a basin wildrye-western grassland and an ecotone of (between) that grassland community and the wheatgrass-big sagebrush savanna.

All of the range vegetation seen here (and in subsequent slides) was on a general, gradual depression (overall, a gently sloping swale) within an overall relatively level land form.

Carter County, Montana. Mid-June; late vernal aspect, soft dough- to grain-ripe stage of western wheatgrass. FRES No. 38 (Plains Grassland Ecosystem), K-57 (Gamagrass-Needlegrass-Wheatgrass). Variant of SRM 607 (Wheatgrass-Needlegrass). Cold Temperate Grassland 142, Plains Grassland 142.1, but no specific Series unit in Brown et al. (1998, p. 40). Northwestern Great Plains- Montana Central Grasslands Ecoregion 43n (Woods et al., 2002).

120. Principal pardners- "Photoplot" of a wheatgrass (western wheatgrass, dominant, thickspike wheatgrass, much less abundant)-Wyoming big sagebrush savanna on a Dense Clay range site in sedimentary Northern Great Plains. This shorter camera-focal distance presented greatr details of botanical composition and structure of this turf(sod-forming)grass-shrub community. The vast proportion of herbaceous phytomass (weight of living plant materior or plant biomass) was of western wheatgrass with that of thickspike wheatgrass substantially less.

121. Escorted by grass- Single plant of Wyoming big sagebrush within a sward of densely rhizomatous western wheatgrass (and small amounts, coverage- and mass-wise, of thickspike wheatgrass). For practical descriptive purposes there were no other grass species in the wheatgrass-big sagebrush savanna treated here. (There must always be an assumption of a few plants of Japanese brome and Kentucky bluegrass on any local disturbance (eg. along a cattle path).

122. Two distinct stands- Contiguous or conterminous with (adjacent to) each other were two distinctly different range plant communities on a slowly permeable (habitat (Dense Clay range site) of overall mixed prairie in the sedimentary Northern Great Plains. Edge of a local stand of western wheatgrass within a wheatgrass-big sagebrush savanna (covered immediately above) in foreground and edge of a basin wildrye (dominant)-western wheatgrass (associate) grassland in background.

Large patch-like areas of these two plant communities along with a transition zone (ecotone) between them (plus local, ephemeral wetlands) formed a vegetational mosaic of Northern Mixed Prairie on a large, gentlly sloping swale in the generally level land form of the unglaciated portion of the Northern Plains.

123. Patches and edges- Mosaic of local populations of basin wildrye (tall, cespitose plants) and western wheatgrass (low-turf or short sodded areas) made up a grassland of these two species which, along with a wheatgrass-big sagebrush savanna and an ecotone of these two communities, formed a landscape pattern of Northern Mixed Prairie.

This pair of photographs provided a "nested photoplot" perspective of the basin wildrye-western wheatgrass community. The simple composition and struction of this range vegetation made for an amazingly bountiful (and beautiful) grassland and the basis of a productive range ecosystem. The pronounced two-layer feature of this grassland was distinct in the second photograph. A wider view of these same features was also provided in two two-slide/caption units below.

Carter County, Montana. Mid-June; late vernal, soft dough- to aspect grain-ripe stage of western wheatgrass. FRES No. 38 (Plains Grassland Ecosystem), K-57 (Gamagrass-Needlegrass-Wheatgrass). Variant of SRM 607 (Wheatgrass-Needlegrass). Cold Temperate Grassland 142, Plains Grassland 142.1, but no specific Series unit in Brown et al. (1998, p. 40). Northwestern Great Plains- Montana Central Grasslands Ecoregion 43n (Woods et al., 2002).

124. Standing tall in the moist north- A beautiful population of basin wildrye dominanted this patch of Northern Mixed Prairie on which western wheatgrass was the associate species. This basin wildrye-western wheatgrass grassland developed on a gradual, gently sloping swale on the overall level surface of the sedimentary Northern Great Plains. Basin wildrye was able to thrive due to extra soil moisture available on this Dense Clay range site.

125. Enter a wetland community- An ephemeral wetland formed as a small pond within 1) surrounding wheatgrass savanna, 2) basin wildrye-western wheatgrass grassland, and an ecotone of these to complete the communities that comprised the landscape of Northern Mixed Prairie in the sedimentary Northern Great Plains. The vegetation of this wet-weather wetland consisted of local colonies of pale spikerush (Eleocharis macrostachya).

The simple structure and species composition of the surrounding basin wildrye-western wheatgrass community evident in these two slides. The tall bunchgrass plants were those of the cespitose (tufted) basin wildrye while the lower-growing sod was maade up of the strongly rhizomatous western wheatgrass. Surprisingly enough there were almost no plants of other species other than a few isolated groups of Japanese chess, an almost always present naturalized Eurasian annual, and an occasional spot of the native, perennial sodformer, inland saltgrass. Both of these grasses were so limited as hardly merit mention.

Carter County, Montana. Mid-June; late vernal aspect grain-ripe stage of western wheatgrass. FRES No. 38 (Plains Grassland Ecosystem), K-57 (Gamagrass-Needlegrass-Wheatgrass). Wet to semi-wet variant of SRM 607 (Wheatgrass-Needlegrass). Cold Temperate Grassland 142, Plains Grassland 142.1, but no specific Series unit in Brown et al. (1998, p. 40). Dense Clay range site (Ross and Hunter, 1976, p. 13). Northwestern Great Plains- Montana Central Grasslands Ecoregion 43n (Woods et al., 2002).

126. An aquatic community added to the mosaic- A wet-weather wetland (ephemeral pond) supported local populations of pale spikerush so as to comprise a fourth community in the landscape of this Dense Clay range site in the unglaciated Northern Great Plains in southeastern Montana. Around perimeter of the ephemeral lake the range vegetation consisted of a basin wildrye-western wheatgrass community of mixed prairie. Tall bunchgrasses were basin wildrye whereas the lower-stature sod areas were plants (or populations) of western wheatgrass.

Carter County, Montana. Mid-June; late vernal aspect, soft dough- to grain-ripe stage of western wheatgrass. FRES No. 38 (Plains Grassland Ecosystem), K-57 (Gamagrass-Needlegrass-Wheatgrass). Wet to semi-wet variant of SRM 607 (Wheatgrass-Needlegrass). Cold Temperate Grassland 142, Plains Grassland 142.1, but no specific Series unit in Brown et al. (1998, p. 40). Northwestern Great Plains- Montana Central Grasslands Ecoregion 43n (Woods et al., 2002).

127. Another composite view- Composition and structure of a basin wildrye-western wheatgrass moist grassland that developed around the perimeter of an ephemeral wetland on a Dense Clay range site in the Northern Great Plains. Domination of this range plant community rocked back and forth between the tall, bunchgrass, basin wildrye, and the rhizomatous, sod-forming western wheatgrass. The areas with visible bare soil were populated mostly by smaller (younger?) plants of western wheatgrass with a very few small spots of inland saltgrass.

Western Wheatgrass True Prairie

The following short section was duplicated in the chapter, entitled True Prairie. While that was the most accurte designation the admittedly arbitrary designation between true prairie and mixed prairie where these two major associations of Great Plains grasslands converge and blend is such that such duplication was deemed desirable in attempting to avoid reader confusion (to the extent possible). Western wheatgrass (Agropyron smithii) is one of the most widely distributed midgrass species over much of the Great Plains. Its biological (species) range extends from (Barkworth et al., ). Western wheatgrass is also a species with almost as wide a distribution of scientific names (binomials). It was explained in the Names section of Introduction of Range Types that over course of the last quarter the genus of western wheatgrass century had changed numerous times including Elytrigia, Pascopyrum, and Elymus (besides Agropyron).

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

133. Purty sparse- Western wheatgrass consociation in the Big Horn Basin portion of the Wyoming Basin in southcentral Montana that could be viewed as either mixed or true prairie though best regarded as mixed prairie as it was nearly an arid precipittion zone, and certainly a xeric edaphic habitat. There were a few plants of Wyoming big sagebrush (eg. left midground of firest slide) and some plains pricklypear and blue grama (as shown in the immediately succeeding slide), but this was essentially a single-species stand, a consociation, of western wheatgrass with needle-and-thread as the dominant and associate species, respectively, the climax decreasers for this dense clay, saline range site.

This is an example of about the most semiarid (xeric soil) environment in which western wheatgrass can survive. This country was in a mild drought, but the herbage production (yield of grass biomass) and cover of western wheatgrass is typically about as low as possible for this cool-season species, this rhizomatous midgrass, to survive.

Carbon County, Montana. Late June-early estival aspect. FRES No. 38 (Plains Grassland Ecosystem. K- 59 (Wheatgrass-Needlegrass), SRM 610 (Wheatgrass). Closest biotic community in Brown et al. (1998, p. 40) was Cold Temperate Grassland 142, Great Basin Shrub-Grassland 142.2, Wheatgrass Series 142.21, and that unit was not it as this western wheatgrass-dominated grassland was not in the Great Basin. Range site was Western Sedimentary Plains- Dense Clay-Clayey-Saline Upland Complex, 5-9 inch precipitation zone (Ross and Hunter, 1976, p. 20-21). Wyoming Basin- Bighorn Basin ecoregion 18b (Woods et al., 2002).

134. Purty rough- Plains pricklypear on a consociation of western wheatgrass with slight cover of blue grama in the Wyoming Basin (specifically, Big Horn) in southcentral Montana. These range plants were quite drought-stressed, but these clayey, saline soils are droughty under best of soil moisture conditions.

Location note: the climax range vegetation of the Wyoming Basin and adjacent areas is a mosaic of natural plant communities ranging from desert scrubland to shrub-grass savannahs to extremely xeric grasslands that could be regarded as seimdesert grasslands. Contiguous with the semiarid western wheatgrass-dominated grassland shown here was a salt desert scrub of Gardner's or Nuttall's saltbush (Atriplex gardneri= A. nuttali). This salt desert scrub or Nuttall's saltbush desert was shown in Range Types of North America under the Shrubland chapter entitled Miscellaneous Shrublands.

The following examples of western wheatgrass plants were taken from the Southern and Central Great Plains. They were included here as the emphasis was on the range plant species and not on the range vegetation such that location from which representative samples were taken was irrelevant. Locations were always given in this publication.

135. Stand of western wheatgrass- Closer-in-view of a consociation of western wheatgrass. This colony may well be many shoots of a single plant of this rhizomatous species. This is another example of a clonal organism in which each shoot is a module, clone or ramet of the genetic individual known as a genet. Peak standing crop: grains are in hard dough stage. Oldham County, Texas. June

136. Standing tall and very thick- Shoots of rhizomatous western whestgrass at peak standing crop. This is a still-yet closer-in-view of the stand shown in the preceding slide. A small (but very sexually repoductive) cheatgrass (Bromus tectorum) plant held its own to clomplete its life cycle in the foreground. This is a good example of sexual vresus asexual reproduction in an annual versus a perennial grass, and of an annual introduced (from Eurasia) invader versus a perennial native decreaeer. Western wheatgrass "hedged its bets" by using both asexual (numerous shoots, modules or ramets, from rhizomes) and sexual (grain production in its many florets as were shown below). Range Management students should familarize themselves with the concept of adaptative "plant strategies" developed by Grime (1979).

Oldham County, Texas. June. Peak standing crop; hard dough stage.

137. Western wheatgrass- An example of western wheatgrass showing the characteristic growth pattern produced by clonal growth and development of "daughter plants" in this rhizomatous species. Lincoln County, Colorado. June.

138. Spike inflorescence of western wheatgrass- The spike arrangement of spikelets is a key tribal characteristic of the Hordeae (= Tritaceae). Erath County, Texas. Mid-May; anthesis.

139. Close-up of spike portions of western wheatgrass- Detail of western wheatgrass spikes showing individual florets in spikelets.Oldham County, Texas. June.

Further study of western wheatgrass:

Spiked on the Northern Great Plains- Three-progressively closer views of spikes of western wheatgrass in anthesis that were growing in the Red River Valley (part of the Central Lowlands physiographic province) in southeastern North Dakota.

Cass County, North Dakota. Late June; peak anthesis.

Beauty in a grass flower- Three views of flowering (specifically, anthesis) in spikes of western wheatgrass growing in the Red River Valley in southeast North Dakota, part of the vast Central Lowlands physiographic province. When western wheatgrass is at this stage of flowering (anthesis) is becomes very showy and strikingly beautiful (even if rather subdued). Yes North Dakota, grasses are flowering and fruit-bearing plants (angiosperms) which, of course, most folks in the Peace Garden State know given that they see it all around them.

And nothing would due but to choose as the North Dakota State Grass (what else?), western wheatgrass. Western wheatgrass is also the state grass of South Dakota and Wyoming. (Some states get some things right.)

Cass County, North Dakota. Late June, peak bloom stage of phenology.

140. Dog town on the state grass- Town of black-tailed prairie dog on a mixed prairie range that is a consociation of western wheatgrass. This range had been and was being heavily grazed--in fact, overused-- by these rodents. Overuse (= overutilization) is defoliation by grazing that exceeds the proper degree of use (greater than proper use factor). Heavy grazing and overuse are NOT synonymous. Under certain management situations and ecological habitats heavy grazing (like light grazing or moderate grazing) can be proper grazing (= proper use). If overuse (excessive defoliation by grazing animals) continues for a long enough period of time (or at critical seasons, phenological stages, and/or under times of stress like drought) changes occur in the plant range community. The more palatable plant species (which are usually the higher seral or climax species) gradually die out because they cannot survive with foliage removal that exceeds what is proper (= sustainable) use for them, plus they cannot compete with plant species that can sustain heavier defoliation or that are not being defoliated. As the more palatable--and usually higher successional order--species die they are replaced by less palatable plant species that are not being grazied as heavily. This causes changes in the range plant community as some plant species replace other range species. This is the state called overgrazing. Overgrazing is prolonged overuse that leads to changes in the range plant community with species of lower succsssional state replacing plant species of higher succcessional order. This phenomenon is retrogression on the sere. Retrogression is the process of range deterioration. Perhaps more precisely stated, range deterioration is the outcome of the process of retrogression.

Composition of the range plant community is measured and known as range condition class. Range condition (class) is the successional state of the range vegetation at a point in time. Changes in in range condition over time is range trend. Downward range trend is the measurement or measured result of range retrogression. With continued (prolonged) overuse the point of overgrazing is reached so that range retrogression (range deterioration) is obvious. . Overgrazing is long-term overuse. Actually overgrazing is the management practice (mismanagement) that leads to changes in range condition class, two or more measures of which indicate range trend. In successional terms retrogression is the retrograde direction of change down the range sere. Retrogression is the reverse of plant succession (though not usually the exact opposite series of changes). Upward range trend is measured (the measurement of) continuing plant succession. This is range improvement via natural development of range vegetation. It is progression upward along the range sere toward climax or, at least, a higher seral stage than was measured the last time range condition class was determined.

A restatement of the phenomenon of range plant succession, of vegetation development, in relation to degradation of the range on this prairie dog town is the process of retrogression or decline of the range plant community down ther range sere. Retrograde movement down the range sere is range deterioration or degradation. A degraded range is one that is undergoing retrogression. Loss or the desired range plant community (= the range plant composition chosen by management) to an extreme degree is range depletion, the result of which is a depleted range.

Before--usually long before--the state of range retrogression commences (or at least is readily detected) there are undesirable changes in individual plants of existing plant species. Variables such as plant size, plant color, number of shoots or leaves, success of sexual reproduction, etc. constitute overall plant health which is known as plant vigor. There are notable changes in plant vigor (decline in general plant health and robustness) that occur with overuse long before the point of overgrazing is reached and retrogression takes place. The astute range manager is alert to such undesirable changes subtle though they are. This is the ability to "read the range". Losses of vigor in plants of desirable (climax or high seral stage) range species are some of Nature's early warning signs of impending decline in range condition and downward range trend.

It was decline of vigor in plants of western wheatgrass that was visible in these two photographs. The western wheatgrass range seen here was being overused. Continued excessive defoliation (by black-tailed prairie dogs in this case) was weaking plants of western wheatgrass. However, at the point of overuse shown in these two photographs excessive defoliation had not reached the state of overgrazing and range depletion. Stay tuned.

Plate County, Wyoming. Late June (early estival aspect). FRES No. 38 (Plains Grassland Ecosystem). K-59 (Wheatgrass-Needlegrass) SRM 610 (Wheatgrass). Closest description in Brown et al. (1998, p. 40) was Cold Temperate Grassland 142, Plains Grassland 142.1, Mixed "Short-Grass" Series 142.13, but this missed more than hit it; Wheatgrass Series 142.21 of Great Basin Shrub-Grassland picked up the wheatgrass, but this was not in the Great Basin and it was the wrong species of wheatgrass. High Plains- Flat to Rolling Plains Ecoregion, 25d (Chapman et al., 2006).

141. Doggone turf- Actually it is the turf more than the prairie dogs that is gone. This was a closer-in "photoplot" of the overused western wheatgrass range introduced in the preceding two slides. Overuse of western wheatgrass on mixed prairie range by black-tailed prairie dogs had exceeded proper use for this range plant species. This sparsely covered, broken turf of western wheatgrass, a consociation of this cool-season grass, was the result of prolonged overuse by prairie dogs. This appeared to have reduced competition (at least for soil mositure and light) between the festucoid grass and plains pricklypear, this latter plant being the asssociate species of this range vegetation. With decline in vigor, western wheatgrass was becoming less effective in competing with plains pricklypear. Tthe result was that the cactus was increasing in both absolute cover and relative cover compared to western wheatgrass.

This obvious shift in overwhelming dominance by western wheatgrass to a situation of greatly reduced abundance of grass tillers (much less cover and denstiy of shoots) and a commensurate increase in cover, density, etc. of plains pricklypear was evidence (early symptoms) of the onset or impending onset of overgrazing by a native range herbivore. Though this relative change in measurable, observable effects was obvious it was still a subtle change because there had not been a relative change in position of dominant and associate range plant species. Furthermore, at this point of severe overuse and onset of overgrazing there was not yet much cover or establishment of other invading species. A few plants of Kochia scoparia (a naturalized, annual, Eurasian forb) were present, but only at trace (or somewhat higher) proportions.

Overuse had just reached overgrazing or maybe it was still closely approaching the point of overgrazing, but changes in plant vigor were glaring in the rangeman's eyes. Even at this "late date" (this state of change in relative plant cover and decline in vigor of the climax decreaser) there was still time to avert range degradation. Reduction in numbers of prairie dogs (such as poisoning part of the prairie dog population) could halt the spread of damage to the range. At least size of area being adversely affected could be stabilized (kept to that part of the range already impacted). Application of poison bait on the perimeter of the town could be used to halt the advance of range decline.

Plate County, Wyoming. late June (early estival aspect). FRES No. 38 (Plains Grassland Ecosystem). K-59 (Wheatgrass-Needlegrass) SRM 610 (Wheatgrass). Closest description in Brown et al. (1998, p. 40) was Cold Temperate Grassland 142, Plains Grassland 142.1, Mixed "Short-Grass" Series 142.13, but this missed more than hit it; Wheatgrass Series 142.21 of Great Basin Shrub-Grassland picked up the wheatgrass, but this was not in the Great Basin and it was the wrong species of wheatgrass. High Plains- Flat to Rolling Plains Ecoregion, 25d

142. Doggone grass; lower on the sere- State of overgrazing on a black-tailed prairie dog town on a mixed prarie range that had been a consociation of western wheatgrass. This was part of the same range (same pasture or grazing unit) that was introduced and described in the two immediately preceding photograph-caption sets. The part of the range shown in this slide was one of the older portions of the prairie dog town. Black-tailed prairie dogs had colonized (established themselves on) this part of the range much earlier than the parts shown above which were outskirts of recent settlement. Over passage of time heavy grazing by prairie dogs had so weakened a once-dense stand of western wheatgrass, the climax dominant grass, that this dominant decreaser had been killed out and replaced by Sandberg('s) bluegrass (all the little bunches of brown grass). Sandberg's bluegrass (a native, cool-season perennial) had completed its annual cycle and had entered dormancy. On this range site Sandberg bluegrass is an increaser.

This impact on western wheatgrass was consistient with that reported by researchers for Great Plains mixed prairie. Agnew etal. (1986) found that grazing, clipping, and burrowing by black-tailed prairie dogs decreased western wheatgrass and blue grama but increased cover of buffalograss and forbs. Uresk (1985) also reported decreased production of western wheatgrass on towns of black-tailed prairie dogs.

At the stage of overgrazing (of retrogrsssion; of range degradation) on this consociation of western wheatgrass Kochia scoparia and buffalobur (Solanum rostratum) had established growing populations which could be seen as short, gray-green or bluish-green little clumps of plants. Almost all western wheatgrass had been eliminated by overuse (prolongued, excessive defoliation) by prairie dogs. Fringed sage (Artemisia frigida) had also made its appearance on the deteriorating range, mostly on the edges of entrances to prairie dog burrows (center mid-ground of photograph). Cheatgrass had not established on this range other than perhaps an isolated rare plant.

Except for K. scoparia all species are native. Black-tailed prairie dog and North American buffalo (Bison bison)--perhaps but not necessarily in that order--have been the dominant range mammals of mixed prairie range for millenia. Temporal and spatial dimensions and population dynamics of these two native mammals (and their forage plants) on the central grasslands of North America prior to human entry by the red man or even before establishment of first white men is known only to God. The time of initial occupation by PaleoIndians (North American Indians) remains a highly debated topic with estimates ranging from as many as 40,000 years to as few as 8,000 years before present. What influence(s), if any of substance, American Indians had on native fauna and, in this case, on black-tailed prairie dogs will likely never be known (at least not in this world). It appears to be self-evident that Indians had to have had some impact on North American buffalo as they successfully hunted them (and all other species of major vertebrates). It must be remembered, however, that some of the more efficient hunting of buffalo by the red man was possible only after mastery of horsemanship that made possible by the white man.'s introduction of the horse. Horseback hunting of the plains buffalo came after mass exterminations of Indians by diseases (smallpox and ubonic plague being the big ones) introduced into aboriginal populations, also by the white man.

Is it possible that prior to arrival of Europeans American Indians had exterminated some animal (and plant) species, the absence of which could have had influences on black-tailed prairie dogs? This is another fiercely debated topic. For instance, could Indians have exterminated predators of prairie dogs? What impacts, if any, did diseases like brucellosis (Bang's disease) and Bubonic Plague, which were introduced by the white man, have on--directly or indirectly--prairie dog populations. For example, canine distemper has had major detrimental effects of the endangered black-footed ferret (Mustela nigripes), one of the major predators of prairie dogs. The black-footed ferret is so susceptible to canine distemper that this disease almost finished exterminating the last black-footed ferrets living in the wild. Guess how canine distemper arrived in North America.

The most fundamental question is, Did the black-tailed prairie dog overgraze the range under completely natural conditions? Was there on-going patch dynamics (retrogression and subsequent secondary plant succession) due to prairie dog grazing before human influence? Did arrival of non-native humans alter the role and/or effect of prairie dogs on the range vegetation of the North American grasslands?

What is now known is that large towns of black-tailed prairie dogs can have major detrimental impacts through overgrazing on mixed prairie rangeland types.

Organiztion Note: another example of overgrazing by black-tailed prairie dogs was presented below in this chapter in the section entitled Shortgrass Prairie Sampler and, also, in the Nebraska Sandhills section of the chapter, Tallgrass Prairie- Interior II.

North American buffalo was adopted as the State Mammal of Wyoming on 23 February, 1985. Western wheatgrass was adopted as the State Grass of Wyoming on 28 February, 2007. OK, so it was eight years after the Wyoming legislature honored a mammal that it got around to honoring a major grass species on which the state mammal largely depends. In spite of this "cart in front of the ox" arrangement (the typical human perspective in regard to plant and animal symbiosis) this great range state did select the species that are most representative of much of flora and fauna of the original grasslands within the Equality State. On the other hand, maybe the black-tailed prairie dog should be the State Mammal of Wyoming. Which one of these two herbivors has (had) the greater influence on the grassland ecosystem? Which is the dominant? Are buffalo and prairie dog co-dominants?

One thing is certain.The rangeman, manipulator of range ecosystems, has the option of managing the range for results like this shown here or in a higher state of range condition. It is up to the range manager, not the prairie dogs.

Note on location and organization: numerous other examples of western grass-dominated grasslands (including western wheatgrass consociations) were presented in the chapter, Mixed Prairie-- Southern and Central Great Plains.

Ecological clarification and note on location/organization: Range types of Northern Great Plains grasslands, including most those of the Northern Rocky Mountain foothills on the east side, have traditionally been interpreted as true prairie (Clements, 1920, ps 121-131; Weaver and Clements, 1938, ps. 518-520) not as mixed prairie. These true prairie range types commonly have a shortgrass component usually including blue grama and, less often, buffalograss. Although shortgrass species, especially blue grama, are typically present they have not been regarded as comprising adequate cover so as to form a vegetational layer, at least not consistently. Similarily, there are occasional tallgrass species in these Northern Great Plains grasslands with substantial cover at local scale, but this cover, frequency, and general abundance are not sufficient to form a distinct tallgrass layer. Furthermore, little bluestem (Andropogon scoparius), the major or most common and abundant of species that attain tallgrass heights to the south and east, thrives as a midgrass species in the Northern Great Plains where it is no taller (and, frequently, shorter) than such midgrasses as sideoats grama and western wheatgrass.

Given this physiogonomy, structure, and species composition most climax grasslands of the Northern Great Plains historically have been described as true prairie. That convention was followed herein. For coverage of major grassland range types of the Northern Great Pains Region viewers were directed to the chapter, True Prairie, Grasslands.
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