Tallgrass Prairie (Interior)-II

Semiarid and Postclimax

The tallgrass prairie includes range vegetation that can be visualized as consisting of two major forms: 1) once-vast grasslands in the continental interior of North America and 2) zones of grasslands and marshes along the shores of North America extending from the Atlantic Coast through the Gulf of Mexico. Climate of the former is continental; that of the latter is maritime. Species composition (including dominants) of plants and animals is similar (sometimes nearly the same) on both of these two basic forms or expressions of tallgrass range. Differences in species in the pre-Columbian ecosystems of these forms was probably not great as there were similar species (= similar ecological niches) or ecological equivalents among range types in them. For example, the dominant climax gallinaceous birds of the interior tallgrass bluestem prairies (eg. Flint Hills and Osage Questas of Kansas and Oklahoma), Gulf of Mexico coastal cordgrass prairies and marshes, and small patches of northern cordgrass prairies along the Atlanic Coast were one subspecies of the grouse known as greater prairie chicken (Tympanuchus cupido pinnatus), Attwater prairie chicken (T. cupido attwateri), and heath hen (T. cupido cupido), respectively. Of course the heath hen is extinct, the Attwater in dire trouble, and the greater is declining, but such similarity in taxa of the major grassland bird of each of these recognized generic grasslands illustrated the ecological relatedness of these major forms of tallgrass grasslands and various range cover types therein.

For pedagogic purposes and convenience (ease of viewing examples) the tallgrass prairie sub-formation (or sub-biome) of North Amereican grasslands was divided into two separate chapters designated as interior tallgrass prairie and coastal tallgrass prairie. There are marshes (herbaceous plant communities usually with the soil surface covered by standing water most of the year or growing season) in (or affilitated with) both of these major forms of tallgrass prairie. Marshes have often interpreded as different from grasslands, perhaps even as separate biomes. Natural distinction between prairie (dominant plants are species of grasses hence grassland) and marsh (dominant plants include species of grasslike plants such as sedges, bulrushes or tules, rushes, cattails, etc. as well as grasses). Designations and differences are often unclear or even arbitrary as, for example, distinction between wet prairie and marsh. The same dominant and associate species of plants are sometimes common to adjacent grassland and marsh. This condition is more common in coastal prairies and marshes than in interior prairies and associated wetlands. Generally marshes are more common and prominent plant communities in coastal than in interior grassland vegetation. Such was reflected in names of these two major forms of tallgrass prairie.

It could be argued that the term prairie could or should encompass both grassland and marsh thereby uniting these two general units of range vegetation. Perhaps prairie could be interpreted so as to include both grassland and marsh. Such a union is not possible given the traditional designation of vegetational or land resource area 2 in Texas as Coastal Prairies and Marshes. The long-standing convention of this "purple-pedigreed' title was retained in the current publication. Rangemen simply have to learn to live with a certain amount of ambiguity.

In the Gulf Prairies and Marshes there are range plant and animal communities that are not strictly speaking dominated by tallgrass species. These include such habitats as seashores and salt flats. Such range vegetation or plant and animal communities are, however, part of the Gulf Prairies and Marshes landscape or greater ecosystem (when seen from pserpective of Landscape Ecology or Ecosystem Ecology). In context of a publication devoted to range types these units of range vegetation were distinctive and different enough from described rangeland cover types (Shiflet, 1994)-- and obviously essential or integral to development and functioning of recognized range types-- that such vegetation was included herein.

The ultimate source for native vegetation of the tallgrass prairie (and true prairie) is North American Prairie (Weaver, 1954). Definitive reference for tallgrass prairie from an ecosystem perspective is that of Kucera (in Coupland, 1992, ps. 227--268).

Nebraska Sandhills (and related postclimax tallgrass types)

Sand and sandy soils (also gravel for that matter) have a higher proportion of macropores (noncapillary pores) so that there is less water runoff and more water that infiltrates this rooting material for plants. Conversely,there is less water retained in the coarse or heavy textured soils as compared to fine (light)-textured soils such as clay; however, , more of the retained water in coarse textures (eg. sand) is available for plant absorption. Also many plants that are adapted to coarse-textured soils have a high proportion of their roots near the land surface so that they absorb this temporarily shallow water before it evaporates or percolates below their root zones. Net result is that the native vegetation on sand, sand dunes, and high sand-content soils is typically more mesophytic than vegetation on adjacent or neighboring soils comprised of less coarse particles (those that have finner soil texture).

This was explained in Weaver and Clements (1938, p. 203-204) by the Clementsian concept of chresard. Chresard is that part of total soil water, the whole content of water in soil (= holard), that is available for plant absorption or use by the plant part. The echard (to withold) is that portion of water still present (held or retained) in soil at the permanent wilting point (ie. water that is unavailable for plant use; water that adheres to tightly to soil particles that plants cannot absorb it). Holard minus chresard equals echard (or any other arrangement of these portions ssuch as holard minus echard equals chresarde or chresard plus echard equals holard). In the Clementsian monoclimax model natural vegetation of sandy environments if postclimax "In Nebraska the relation [transition from true to mixed prairie] is further disturbed by the extensive sand-hill region, in which the high chresard favors a postclimax of tall grasses far beyond their proper climate [ie. beyond the climatic or regional climax]" (Weaver and Clements, 1938, p. 524). "In the vast sand-hill area of Nebraska, the tall-grass postclimax attains its best development, which is assumed to reflect the climate when the prairies were occupied by the bluestems and their associates some millions of years ago" (Weaver and Clements, 1938, p. 86). This also included shrubs such as New Jersey tea, sand sagebrush (Artemisia filifolia), and soapweed yucca.

The sandhill region of Nebraska and similar smaller areas in other parts of the Great Plains including some in Colorado, Kansas, Oklahoma, Texas, and New Mexico support more mesic (more moisture-requiring) range vegetation than that of less porous soils in the same region or area. In the vernacular of Tansley's polyclimax model or Whittaker's climax pattern interpretation postclimax vegetation would be edaphic and/or topographic climax vegetation (edapho-topographic climaxes). Same ecological (successional or climax) outcome.

Tallgrass prairie range vegetation in the Nebraska Sandhills (and some similar range communities in Colorado, Oklahoma, New Mexico, Texas, etc.) is in the Mixed Prairie Region though, of course, it is not mixed prairie. As such, postclimax or edaphic-topographic climax tallgrass prairie was covered in this chapter. This included such range types as sand sagebrush- and shinnery oak (Quercus havardii)-tallgrass savannas as well as the classic Nebraska Sandhills tallgrass prairie. Range types of mixed prairie, including some in the Nebraska Sandhills (not all Sandhills grassland is tallgrass prairie), were (logically) dealt with in the chapter, Mixed Prairie.

The Nebraska Sandhills also have wonderfuly fascinating wetlands including marshes and wet prairie (sometimes complete with local stands of cottonwood and willow). Examples of his range vegetation were also included in this section although marshes and range vegetation dominated by woody species are not, strictly speaking, grassland. This was explained in greater introductory detail above under the section entitled, Wet Prairies and Marshes of Tallgrass Prairie.

Organization Note: All (or almost all) of the famed Nebraska Sandhills are in the semiarid zone. This is mixed prairie country, part of the Mixed Prairie (Stipa-Bouteloua Association) of Clements (1920, ps. 135-139). Tallgrass prairie has developed on that part of the eastern Sandhills of Nebraska which through a combination of deeper, sandier soil and slightly higher annual precipitation is a postclimatic environment (hence, tallgrass prairie in the Nebraska Sandhills is postclimax vegetation). The rest (western portion) of the Nebraska Sandhills is Mixed Prairie (the true climax, the climatic climax of the region). Mixed Prairie in the Nebraska Sandhills was treated under the Mixed Prairie chapter.

1. A warty hided "Howdy" from the Nebraska Sandhills- A Great Plains toad (Bufo cognatus) near prairie rose (Rosa arkansana) in the heart of the Nebraska Sandhills seemed an appropriate vertebrate greeter to this grassland Garden of Eden. This amphibian can readily burrow through the loose (and (sometimes moist) sandy soil that characterizes the Sandhills. Presence of Great Plains toad thus served as a cold-blooded barometer of the soil component of this range ecosystem.

At large mapping scale the major soil area over much of the Nebraska Sandhills is Valentine-Nueces-Dune sand (Sands, dry). This soil is an example of an azonal soil, Azonal order or category VI (United States Department of Agriculture, 1938, ps. 995, 1136-1137). This azonal soil group is relatively restricted in land area in contrast to the more ridely distributed zonal or intrazonal soils. Likewise, parent material (eolian sand in this instance) was the major determining soil formation factor. Currently the Valentine soil series is much more restricted by the system of Soil Taxonomy whereby it was described as mixed, mesic typic ustipsamments. There is a Valentine soil association as well as several associations in which Valentine is a major, defining soil (various Nebraska county soil surveys--if you can find them, and good luck).

The Great Plains toad is a widely distributed amphibian with a species range extendingh from the Canadian Prairie Provinces to central Mexico. A detailed monographic treatment of this range toad was that of Diduk (1999, updated generically 2002). The toad was not what most of us think of as a range animal though of course it is. Nice way to welcome rangemen to the Nebarska Sandhills.

Thomas County, Nebraska. Mid-June.

2. Farmed Sand Hills of Nebraska ("God's Own Cow Country")- This western-most extension of tallgrass prairie is typically a community of the Four Horsemen species. Region-wide little bluestem, State Grass of Nebraska, is clearly dominant in an abiotic environment more characteristic of tall and true prairies than the mixed prairie that would be expected in this semiarid precipitation zone. The Sand Hills range vegetation type is the product of soils of deep sand, often accompanied by shallow surface acquifers. In classic Clementsian view this tallgrass prairie is postclimax to both true and mixed prairie. This is a Choppy Sands range site with sand bluestem (Andropogon hallii), sand lovegrass (Eragrostis trichodes), and prairie sandreed (Calamagrostis longifolia) the dominant species. Cherry County Nebraska. Hiemal aspect, October. FRES No. 39 (Prairie Ecosystem). K-67 (Nebraska Sand Hills Prairie). SRM 602 (Bluestem- Prairie Sandreed). Nebraska Sand Hills- Sand Hills Ecoregion, 44a (Chapman et al., 2001).

3. Sands range site of the Nebraska Sand Hills- Same species as above with more mid and short grasses like blue and hairy grama (Bouteloua gracilis,B. hirsuta) and Junegrass. Cherry County, Nebraska. Hiemal aspect, October. FRES No. 39 (Prairie Ecosystem). K-67 (Nebraska Sand Hills Prairie). SRM 602 (Bluestem- Prairie Sandreed). Nebraska Sand Hills- Sand Hills Ecoregion, 44a (Chapman et al., 2001).

4. Sandhills Marsh- Wetland range site.Mostly grass-like plants such as rushes (Juncus spp.), cattail (Typha angustifolia, T. latifolia), bur-reed (Sparganiumeurycarpum ) locallywith sedges (Carex spp.) throughout.Note lodge of muskrat (Ondrata zibethica). Cherry County Nebraska. As part of tallgrass prairie this is in FRES No. 39, but as a marsh it could as logically be placed in the tremendously varied FRES No. 41 (Wet Grassland Ecosystem). Either a hydric variant of K-67 (Nebraska Sand Hills Prairie) or K- 42 (Tule Marshes). Wetlands variant of SRM 602 (Bluestem- Prairie Sandreed). Nebraska Sand Hills- Lake Area Ecoregion, 44d (Chapman et al., 2001).

5. Early summer in the Nebraska Sandhills- This slide introduced a series of photographs that showed the sand and/or big bluestem-prairie sandreed-dominated Nebraska Sandhills in early summer coming out of a five-to-six year drought. The ranges chosen for inclusion in this series were, except for some purposely included blowouts, in the climax (or nearly so) state (Excellent range condition class). These pastures either had been or were currently being grazed by cow-calf pairs so that there was considerable standing dead herbage from last year's grass crop.

The range introduced here was a species-rich postclimax tallgrass community with an amazing array of tall-, mid-, and shortgrasses including sand bluestem, upland switchgrass, prairie sandreed, little bluestem, Canada wildrye, sideoats grama, blue grama, and hairy grama as common species. There was also small amounts of sand dropseed, buffalograss, and, in lower moist microhabitats, western wheatgrass.

Cherry County, Nebraska. Late June; early estival aspect. FRES No. 39 (Prairie Ecosystem). K-67 (Nebraska Sand Hills Prairie). SRM 602 (Bluestem- Prairie Sandreed). Bluestem "Tall-Grass" Series, 142.11 of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Sandy range site (NRCS, 2005). Nebraska Sand Hills- Sand Hills Ecoregion, 44a (Chapman et al., 2001).

6. Magazine cover shot- Textbook view of Nebraksa Sandhills postclimax tallgrass prairie dominated (overall) by sand bluestem, little bluestem, and prairie sandreed with locally important major grasses including blowout grass (Redfieldia flexuosa), sandhills muhly or, sometimes, blowout muhly (Muhlenbergia pungens), sand lovegrass, sand dropseed, needle-and-thread, sideoats grama, and blue grama. The dominant shrub was soapweed or soapweed yucca (shown here at peak bloom stage) which gave a savanna (at least a savanna-like) physiogonomy to this Andropogon-Calamovilfa grassland. This grassland vegetation was not, however, anything approaching a savanna is actual terms. The second most important shrub on this picture postcard range was New Jersey tea (more of this low shrub was shown below).

Obviously this range was in Excellent range condition class. It was selected to show the Nebraska Sandhills grassland at its zenith and to uplift outstanding stewardship of range resources. Blowouts are a constant threat to the integrity of grassland vegetation in the Sandhills such that grazing must be judicious. The wind can bring unforgiving devestation to this beautiful and fragile land. There are many opporatunities for mistakes and few occasions for second chances. Grazing management on this range had been superb; this land owner knew and played by Mother Nature's strict rules.

These two cover photographs were still probably not up to standards of National Geographic. Keep trying was deemed a worthy plan.

Cherry County, Nebraska. Late June; early estival aspect. FRES No. 39 (Prairie Ecosystem). K-67 (Nebraska Sand Hills Prairie). SRM 602 (Bluestem- Prairie Sandreed). Bluestem "Tall-Grass" Series, 142.11 of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Sands range site (Natural Resources Conservation Service, 2005). Nebraska Sand Hills- Sand Hills Ecoregion, 44a (Chapman et al., 2001).

7. Shrub or forb: your call- Soapweed or soapweed yucca (Yucca glauca) in the heart of the Nebraska Sandhills. In the Nebraska Sandhills-one of the most fabulous expanses of unbroken virgin grasslands remaining on Earth--there is remarkably little cover and relatively few species of woody plants. The Yucca species do have wood-resembling tissue and some secondary growth of their stems. Gould and Shaw (1983, p. 7) described Yucca species as being "[a]rborescent, woody plants..." that "... do have a cambium in the cortex...". Some of the Yucca species are of such size, and along with their woody or wood-like compositition, as to be recognized as shrubs or even trees. Sargent (1947, ps. 3-28) included eight Yucca species in The Silva of North America, the ultimate encyclopedia of North American trees. Elias (1980, ps. 904-916) also included eight Yucca species in his defintiive treatment. Thus it would seem that Y. glauca would qualify as a shrub which, in fact, it was regarded when Dayton (1931, ps 14-15) included it with browse plants. Others might regard this species as a forb. Unlike the grass group of range plants, the categories of forbs, shrubs, and trees are somewhat open to interpretation.

In this same context, some authors such as Smith (1977, p. 256) have interpreted tha agave group, of which Yucca is a member, as being a subfamily (Agavoideae) of Liliaceae whereas others like Diggs et al.(1999, ps. 1082-1086) and Kaul et al. (2006, ps. 72-73) see the agave taxon as being a separate family, Agavaceae.

Thomas County, Nebraska. Mid-June-peak bloom phenological stage.

8. At their peak in the Sandhills-Two plants of soapweed yucca at peak bloom in the Nebraska Sandhills. Under overgrazing soapweed yucca responds as an invader (yucca leaves are much less palatable than those of most grass species), but as can be seen in this photograph (and two other slides shortly above) Yucca glauca is also abundant on Sandhills grassland in Excellent condition class. There can be several thousand seeds per Yucca fruit which is a loculicidal capsule (see slides below). There are no shortages of soapweed yucca seeds on Nebraska Sandhills grasslands.

Thomas County, Nebraska. Mid-June; full-bloom stage.

9. Floral details of the Agavoideae-Flowers of soapweed yucca presented so as to show details of this lily-like inflorescence. There are four petals in the perfect flower of Yucca species. These petals appear whitish, but they are also have a greenish tinge and, frequently, with areas of purplish tones (Kaul et al., 2006, p. 72). The six tepals (sepals and petals collectively when they cannot be distinguished, when the perianth is not divided into calyx and corolla) were visible in these photographs. Also visible (in last two photographs) were the developing six stamen and the stigma which develops into a six-lobed organ.

The flowers of Yucca species are some of the most palatable plant parts on the range. They are sought out and eaten with such obvious preference by all kinds of grazing animals that it is as if Yucca blooms are the "ice cream and cake" of the range buffet. Presence of Yucca flowers remaining on the range (and, even more so, ripe inedible fruit that indicated even longer persistence of Yucca flowers) is a sure sign that a given range has not been grazed or, at least, not stocked very heavily.

Thomas County, Nebraska. Mid-June

10. Now for the fruits and seeds- Seed-filled capsules (first slide) and interior of empty (seed-shed) capsules (second slide) of soapweed yucca growing on an Excellent condition class range in heart of Nebrask Sandhills. The fruit of Yucca species has traditionally been hard capsule. Yucca glauca has six locks per capsule due to formation of of partitions between rows of ovules (seeds) in each of three carpels; strictly speaking, the single pistil of each flower was formed by fusion of three carpels resulting in female organ of three locules, each of which has a petition between rows of seeds (Kaul et al., 2006, p. 72).

The shiny black and winged seeds are flat and quite thin such that many of them are contained within each lock of the capsule. It is little wonder that there can be so many plants of soapweed yucca on Sandhills grassland.

Thomas County, Nebraska. Mid-June; post-fruit-bearing phenological stage.

11. Slow slide down a dune- Three-slide sequence serving as a "photo-transect" from north-slope at summit (so to speak) of sand dune (first slide) down through midslope to toe of dune ending at base of this dune at left and progressing to the interdunal area at right (second slide), and finally stopping in center of this interdunal area with south-slopes of another dune in mid-to background (third slide). On the north slope (first slide) eastern red cedar (Juniperus virginiana), represented by two still-small palnts, was starting to invade. Cedars were accompanied by western poison ivy (Rhus rydbergii= Toxicodendron ryderbii= T. radicans var. rydbergii) as featured prominently in lower left corner of first slide and foreground of the second and third slides. Snowberry (Symphorcarpus albus) was a third woody species in this tallgrass prairie range plant community.

Little bluestem and sand dropseed were the dominant grasses in the small basin in foreground of first slide. Switchgrass, readily identified (even at this distance) by tall, rank, dead shoots was the most abundant (and most obvious) plant species in the foreground of second slide where it was joined (as just noted) by western poison ivy. Other major grass species in this second section of the "photo-transect" were sand bluestem and prairir sandreed. All three of theser tallgrasses were still short at about the fourth-to sixth-leaf stage. The microsite of interdunal basin was more mesic than even the north slope of the dune which accounted for the more moisture-requiring switchgrass and more sand bluestem and prairie sandreed than on the dune (even if it was a north slope). The third slide-section of the "transect" included a recent, small blowout that was "hairing over". This local blowout had not completely recovered by secondary succession with the resulting state of dominance by blowout grass with its main associate, sand dropseed, but both had the company ( with some to considerable cover) of sand bluestem, switchgrass, and prairie sandreed as well as the conspicuous western poison ivy. There was also some cover of the naturalized Eurasian annual, cheatgrass or downy chess (as is often the case for disturbances).

Cherry County, Nebraska. Late June; early estival aspect. FRES No. 39 (Prairie Ecosystem). K-67 (Nebraska Sand Hills Prairie). SRM 602 (Bluestem- Prairie Sandreed). Bluestem "Tall-Grass" Series, 142.11 of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Sands range site (Natural Resources Conservation Service, 2005). Nebraska Sand Hills- Sand Hills Ecoregion, 44a (Chapman et al., 2001).

12. Another interdunal basin (depression)- A between-the-dunes "bottom" that was larger than the basin featured and described in the preceding three-slide sequence. The range plant community of the depression shown here included an additional, a more interior, zone the range plant community of which was comprised of sand cherry or, sometimes, sand plum (Prunus besseyi= P. pumila var. besseyi) with Carex and Juncus species having greater cover and density. Surrounding this innermost, "circular" (disk) and more mesic range vegetation was an outer "ring-shaped" zone made up of blowout grass, sand bluestem, switchgrass, prairie sandreede as well as the naturalized Eurasina perennial, smooth bromegrass. Stiff sunflowr (Helianthus rigida) was indiscriminate in choice of edaphic and/or moisture gradients and was growing on both of the microenvironments just described.

The sand cherry-caric sedge-rush "inner circle" was shown exclusively in the second of these two photographs. The first photograph showed both local (microhabitat) zonal plant communities of the interdunal range vergetation with the outer vegetational zone of tallgrasses and western poison ivy featured in the foreground.

Cherry County, Nebraska. Late June; early estival aspect. FRES No. 39 (Prairie Ecosystem). K-67 (Nebraska Sand Hills Prairie). SRM 602 (Bluestem- Prairie Sandreed). Bluestem "Tall-Grass" Series, 142.11 of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Sands range site (Natural Resources Conservation Service, 2005). Nebraska Sand Hills- Sand Hills Ecoregion, 44a (Chapman et al., 2001).

13. Looking up a sandhill and, even more, at what's growing on it- "Photo-transect" from base to summit of a large sand dune of a Choppy Sands range site in the Nebraska Sandhills. This was postclimx tallgrass prairie dominated by sand bluestem and prairie sandreed (the typical tallgrass, decreaser dominants on deeper sands) with needle-and-thread (the major, regional, decreaser midgrass) as the associate species. Also abundant to locally dominant on wind-disturbed microsties were Redfieldia flexuosa and Muhlenbergia pungens (scientific names of blowout grass and sandhills muhly, respectively, were repeated to avoid confusion with the common name of "blowout grass" that is, unfortunately, used interchangeably for these two important species). Blue grama, sideoats grama, Canada wildrye, and plains lovegrass were also important species. Sand dropseed and cheatgrass or downy brome were present but infrequent. Soapweed yucca is one of the most widely distributed woody to semiwoody species across the vast Great Plains and it was well-represented on this range that obviously was in Excellent range condition class. By the way is Yucca glauca a short shrub, a subshrub, or a forb? It has been interpreted and described as any and all of these by one authority or another. Most workers, including this writer, have regarded it as a shrub.

This was pristine range vegetation so invaders were as "scarce as hen's teeth". There were local, small, patchlike blowouts that were effectively stabilized by R. flexuosa and M. pungens, the two main sandbinding invaders of such "open sores" in the Nebraska Sandhills. This is a classic example of patch dynamics on localized, naturally disturbed spots in sandhills grassland. Initiation and local spread of denudation by geologic wind erosion has been an on-goining natural feature of this landform and its plant communities for millenia. This ever-present phenomenon maintains sandbinding range plants in the native vegetation and ensures that blowouts do not expand to such size that they cannot be quickly stabilized by such species. In this context R. flexuosa and M. pungens are generally interpreted as increasers, and certainly essential members of the postclimax range vegetation.

A small blowout had been stabilized midslope on this dune (right midground in front of two soapweed yucca). A new blowout had begun just below dune crest (far right background). This new blowout was shown in greater detail in the next photograph.

Cherry County, Nebraska. Late June; early estival aspect. FRES No. 39 (Prairie Ecosystem). K-67 (Nebraska Sand Hills Prairie). SRM 602 (Bluestem- Prairie Sandreed). Bluestem "Tall-Grass" Series, 142.11 of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Choppy Sands range site. (Natural Resources Conservation Service, 2005). Nebraska Sand Hills- Sand Hills Ecoregion, 44a (Chapman et al., 2001).

14. Atop a sandhill- Range vegetation of postclimax tallgrass prairie from the vantage point of roughly half halfway to the crest of a large sand dune of Choppy Sands range site in the Nebraska Sandhills. This was the summit of the big dune introduced in the immediately preceding photograph.Two things about range plant communities in the Sandhills were featured in this large "photo-plot": 1) details of the Andropogon-Calamovilfa-dominated sward in Excellent range condition calss (diagonally from lower right foreground through central and left mid-to background) and 2) start of a blowout (right margin to summit of dune in right background plus left to center foreground).

Climax tallgrass dominants were sand bluestem and prairie sandreed with the climax midgrass, needle-and-thread, the associate species.Other important grasses were Canada or nodding wildrye, blue grama, and, in far left foreground, sandhills or blowout muhly. Soapweed yucca was the only shrub of consequence.

Blowouts are the main form of patch dynamics in the tallgrass prairie on these sand dunes in the Central Great Plains. Plant succession on wind-induced denudation is a natural phenomenon in this range ecosystem. Proper graaing management must have has one of its major goals maintenance of vigerous grass plants that are capable of invading blowouts and stabilizing these patches or gaps in the grassland sward before they grow to such size that wind erosion becomes accelerated rather than geologic erosion and caused range degradation and even collapse of the grazing ecosystem. The processes of blowout formation and stabilization--and the problem posed by blowouts in this landform and rangeland cover type--were dealt with below.

Cherry County, Nebraska. Late June; early estival aspect with the tallgrass dominants still in early growth (roughly four- to eight-leaf stage).

15. Sward of Nebraska Sandhills tallgrass prairie- "Photo-quadrant" of postclimax range vegetation on a choppy Sands range site dominated by sand bluestem and prairie sandreed (both in early growth stages: four-to eight-leaf stages of phenology). Throughout or across this sample of the range plant community needle-and-thread--a climax cool-season, midgrass--was the associate species. Redfieldia flexuosa and Muhlenbergia pungens (binominals used to avoid confusion with two common names of "blowout grass") were also abundant and even locally dominant on small patches of revegetating blowous. Blue grama, sideoats grama, and canada wildrye was also commonly present.

Cherry County, Nebraska. Late June; early estival aspect. FRES No. 39 (Prairie Ecosystem). K-67 (Nebraska Sand Hills Prairie). SRM 602 (Bluestem- Prairie Sandreed). Bluestem "Tall-Grass" Series, 142.11 of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Choppy Sands range site. (Natural Resources Conservation Service, 2005). Nebraska Sand Hills- Sand Hills Ecoregion, 44a (Chapman et al., 2001).

16 Sandhills or sand muhly (Muhlembergia pungens)- This is one of the more important dominant grasses of the Nebraska Sandhills on disturbed range, especially overgrazed and blowout areas. Sandhill muhly-dominated communities constitute a succcessional stage between that pioneerd by the strongly rhizomatous blowout grass (Reldfieldia flexuosa), the dominant perennial grass of blowouts, and the climax bunchgrass community of little bluestem, sand bluestem, prairie sandreed, and needle-and-thread (Weavr and Albertson, 1956, ps. 173-177). Sandhill muhly typically remains an occasional species in the climax bluestem-sandreed-needle--and--thread prairie. Such was the case for the two plants shown here.

Cherry County, Nebraska. Late June; mid-growth phenological stage.

17. At edge of blowout- Creation of a blowout that had expanded to the stage of "already too big" on a Sandy range site in the Nebraska Sandhills. The problem with accelerated soil erosion by the agent of wind and resultant blowouts was described above. The process of revegetation and stabilization of blowouts by means of secondary plant succession was also noted earlier as well as given in greater detail here and in subsequent slides.

This slide presented the point of contact between an apparently expanding blowout and presently undisturbed postclimax tallgrass (Andropogon hallii-Calamovilfa longifolia-dominated) prairie sward in Excellent range condition calss to the immediate right of the enlarging blowout. Other range plants in the undisturbed (so far) grassland included western poison ivy (short, bright-green broadleaf plants at immediate edge of contact, both in and outside of blowout), sandhill muhly, blowout grass (again, Redfieldia flexuosa to avoid confusion with this common name), needle-and-thread (local associate species), sideoats grama, plains lovegrass, sand dropseed, and blue grama.

To the inexperienced passing-through traveler (including this photographer-rangeman) it is difficult to ascertain whether a given blowout is "growing" (expanding) or being successfully invaded, colonized, and in process of being stabilized by range plants at the margins (outer edges) of the blowout. A fairly reliable indicator is if perennial colonizing grasses like Redfieldia flexuosa and Muhlenbergia pungens (increasers) are on the perimeter of (at least portions thereof) the blowout they are almost certainly advancing into the blowout and starting to stabilize it. By contrast, the blowout is still actively encroaching into intact grassland if range plant species at the perimeter are dominant decreasers like sand bluestem, prairie sandreed, and needle-and-thread. Species mixtures of colonizers and decreaser dominants (and associates like needle-and-thread) at points of contact between blowout and intact grassland complicate "reading sign" at the "crime scene". Status of blowouts and integrity of grassland vegetation is more confusing yet--at least to the inexperienced eye--on ranges in Fair where decreaser tallgrasses are typically in "short supply" even without blowouts. Generally there cannot be ranges in Poor range condition class in the extremly sandy and high dune forms of Nebraska Sandhills (eg. Choppy Sands range site) as such states of retrogression will have inadequate plant cover on the soil surface to prevent extreme wind-caused soil erosion (ie. Poor condition range in the sandhills is an immense blowout).

The blowout presently was from all appearances an active (expanding) one. Rhizomatous colonizing grasses were absent from parts of the blowout-intact grassland contact shown here. Instead sand bluestem, prairie sandreed, plains lovegrass, and needle-and-thread were grasses present (precariously so) on the apparently advanced frontline of the blowout.

The Nebraska Sandhills constitute a delicate, fragile land. Good stewardship of sandhills ranges is essential to survival in ranching . The wind is an unforgiving enforcer of wise use management of rangeland in the sandhills landform.Sound grazing management is that which minimizes the natural cycle of denudation and subsequent secondary plant succession; stability or the range ecosystem and ranching firm is maximized when patch dynamics is kept within the natural cycle of dynamic equilibrium (or even less).

Cherry County, Nebraska. Late June; early estival aspect. FRES No. 39 (Prairie Ecosystem). K-67 (Nebraska Sand Hills Prairie). SRM 602 (Bluestem- Prairie Sandreed). Bluestem "Tall-Grass" Series, 142.11 of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Sands range site (Natural Resources Conservation Service, 2005). Nebraska Sand Hills- Sand Hills Ecoregion, 44a (Chapman et al., 2001).

18. On the mend (or hairing over)- A recent blowout on a Sands range site in the Nebraska Sandhills was undergoing secondary plant succession and slowly being stabilized following invasion, ecesis and aggregation (Weavr and Clements, 1938, ps. 3-4, 117-147) of colonizing grasses. Redfieldia flexuosa, the species most commonly known as blowout grass, was the principal range species that had successfully invaded and was in the process of revegetating this blowout As this strongly rhizomatous species was stabilizing this small spot of sandy soil prairie sandreed and sand bluestem were already stationed at the edge of the denuded space poised to complete secondary plant succession on this patch so as to protect this fragile range ecosystem as has been done for countless cycles. These are the two processes of reaction and stabilization in the Clementsian model of "dynamic vegetation" in which plant succession progresses from invasion following denudation back along the sere to climax.

Soapweed yucca (center background) had "done its duty" to hold this coarse yet delicate soil from being carried away and redeposited by the incessant wind, a major abiotic factor in the Nebraska Sandhills landscape and the Great Plains province in general.

Cherry County, Nebraska. Late June; early estival aspect. FRES No. 39 (Prairie Ecosystem). K-67 (Nebraska Sand Hills Prairie). SRM 602 (Bluestem- Prairie Sandreed). Bluestem "Tall-Grass" Series, 142.11 of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Sands range site (Natural Resources Conservation Service, 2005). Nebraska Sand Hills- Sand Hills Ecoregion, 44a (Chapman et al., 2001).

19. Filling in- Closer-in view of blowout grass (Redfieldia flexuosa) colonizing a blowout on a Sands range site in the Nebraska Sandhills. This is the single most important range plant species for early stabilization of newly formed sand dunes and blowouts in this rangeland cover type, SRM 602 (Bluestem- Prairie Sandreed).

Cherry County, Nebraska. Late June.

20. At and along top of another sand dune- Nebraska Sandhills range vegetation just below crest of a sand dune on an east slope. This east slope was a more mesic than average or typical for this landform. The range plant community on this more moist and overall more moderate east slope included some species with somewhat greater soil moisture requirements. The most conspicuous of these in these two "photo-plots" was New Jersey tea presented here at peak bloom in an early estival aspect and plant society. At local spatial scale New Jersey tea was an associate to the dominant postclimax tallgrass species, sand bluestem and prairie sandreed. This floristic relationship (species assemblage) was shown in the first photograph. The second photograph presented at shorter camera focal length a microhabitat (local, small depression slightly below crest of a taller sand dune) comprised primarily of these same three decreaser species.

Further details of this stand of postclimax tallgrass prairie on this same sand dune at midslope (downhill from near dune crest) were presented in the immediately succeeding two-slide and caption segment.

Cherry County, Nebraska. Late June; early estival aspect. FRES No. 39 (Prairie Ecosystem). K-67 (Nebraska Sand Hills Prairie). SRM 602 (Bluestem- Prairie Sandreed). Bluestem "Tall-Grass" Series, 142.11 of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Sands range site (Natural Resources Conservation Service, 2005). Nebraska Sand Hills- Sand Hills Ecoregion, 44a (Chapman et al., 2001).

21. Close-up of range vegetation at mid-slope of a sand dune- Two "photo-plots" of postclimax tallgrass prairie dominated by sand bluestem and prairie sandreed with new Jersey tea as local associate about "half-way up" (or "down)" a taller dune (at roughly mid-height) in the Nebraska Sandhills. These two photographs were taken at shorter camera focal length and downslope from the range vegetation presented in the two photographs and caption immediately preceding this two-slide treatment. These "closer up" perspectives showed the sward of this stand of tallgrass prairie at distances that featured characteristics of plant cover on the soil surface.

Major plant species were sand bluestem and prairie sandreed. Current season--actively growing--shoots on plants of f these two dominant decreasers were at roughly the four to eight leaf stage of phenology. Some of last year's mature shoots of these two tallgrasses were present as dead and weathered stalks. Needle-and-thread, a cool-season midgrass, was the associate grass species. It was present in this stand at peak standing crop and hard-dough to grain-ripe stages.

This range country had been in drought for the previous five to six years, plus the current growing season (up to the sixth day of summer when these photographs were takne) for warm-season species had been wet but cold (considerably below mean ambient and soil temperatures). Consequently, growth of warm-season native grasses had been slower and later than was tyhpical. This was most likely even more the situation for sand bluestem and prairie sandreed, the two dominants, which are late-blooming and the largest grasses on this range site. By contrast, New Jersey tea, the principal shrub in this stand, was in (at) peak bloom. This shrub of the buckthorn family (Rhamnaceae) was present only in the second of these two slides where it was represented by smaller plants as that. New Jersey tea at full-bloom stage was shown in the sets of slides immediately before and after the current set.

Weaver and Clements (1938, p. 86) that it was "[t]he compensatory influence of sand" with the greatly increased water-holding capacity that was responsible for existence of New Jersey tea as a relict species in the Nebraska Sandhills.

Cherry County, Nebraska. Late June; early estival aspect. FRES No. 39 (Prairie Ecosystem). K-67 (Nebraska Sand Hills Prairie). SRM 602 (Bluestem- Prairie Sandreed). Bluestem "Tall-Grass" Series, 142.11 of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Sands range site (Natural Resources Conservation Service, 2005). Nebraska Sand Hills- Sand Hills Ecoregion, 44a (Chapman et al., 2001).

22. At bottom of a sand dune- General view (first photograph) and detail or species composition view (second photograph) of postclimax tallgrass prairie in interdunal area at base of a large sand dune in the Nebraska Sandhills. Local physiography (relief) in areas among dunes often results in swales such as the one shown here in the first photograph. Blowouts (see above) frequently occur in such interdunal areas which could possibly increase or exagerate depth and plane surface of these swales. Sand does have a tendency to roll down from updune slopes such that microtopography is a result of soil deposition from the countervailing erosional impacts of gravity and wind as agents of soil movement. Smaller, even more localized depressions likely result from the same process and pattern of soil erosion. This may result from geologic (natural) and/or accelerated erosion.

The swale presented here was a recent blowout that had been stabilized by Redfieldia flexuosa, the plant to which the common name of blowout grass is most frequently applied. Sand bluestem, prairie sandreed, and sandhill muhly were successfully invading and at the point in plant succession of aggregation (Weaver and Clements, 1938, ). New Jersey tea was also present, but as full-sized plants for this range site which suggested the possibility that it had persisted to some degree in the blowout or perhaps that the blowout had not been particularly deep and/or that it was more tolerant of sand movement than sand bluestem and prairie sandreed (or maybe these plants were just "lucky"). Stiff sunflower was growing in an interrupted semicircular spatial arrangement around New Jersey tea. Soapweed yucca was common on intact grassland around and up to the perimeter of the blowout.

Comparison of this range plant community at base, mid-height (mid-slope), and crest of the same dune showed the consistency in species make-up, physiogonomy, and structure of this duneland vegetation on a range that, except for blowouts, was in Excellent range condition class.

Cherry County, Nebraska. Late June; early estival aspect. FRES No. 39 (Prairie Ecosystem). K-67 (Nebraska Sand Hills Prairie). SRM 602 (Bluestem- Prairie Sandreed). Bluestem "Tall-Grass" Series, 142.11 of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Sands range site (Natural Resources Conservation Service, 2005). Nebraska Sand Hills- Sand Hills Ecoregion, 44a (Chapman et al., 2001).

23. Sandy shrub- Another (and one of the few) shrubs in the Nebraska Sandhills is Bessey's sand cherry (Prunus besseyi= P. pumila var. besseyi). At time of this writing this taxon was recognized in the "official" (contemporary) flora of the Cornhusker State (Kaul et al., 2006) and in Flora of the Great Plains (Great Plains Flora Association: McGregor et al., 1986) as a taxonomic variety of P. pumila. Previously the Nebbraska Sandhill sand cherry was interpreted as a seperate species, P. bessey, including in the Atlas of the Flora of the Great Plains (Great Plains Flora Association: McGregor et al., 1977). This species-level designation had as its specific epithet a commenerative name honoring Dr. Charles E. Bessey, Founding Father of American Plant Ecology at University of Nebraska and the disciplinary "sire" of such immortals as Frederic E.Clements, John E. Weaver, and Roscoe Pound.

It was also Professor Bessey who left this immortal and pertinent quotation: "Football occupies the same relation to education that a bullfight does to farming". It seemed such a shame to demote the uniqueness of this Prunus species which is so adapted to its unique edaphic/climatic habitat (and to thereby denigrate the sacred memory of Dr. Bessey) that the author of Range Types retained the original species-level of distinction. Prunus besseyi was recognized by Fernald (1950, p. 878) as differing from P. pumila by ascending or erect branches and smaller, coriaceous, sharply toothed leaves in the former. Prunus besseyi was the desingation in The Phytogeography of Nebraska, the seminal work on Nebraska vegetation, authored by Pound and Clements (1900) and published by the University of Nebraska Botanical Seminar.

Cherry County, Nebraska. Late June.

24. Sandy (and rosy) shrub a little farther south- Another example of Bessey's sand cherry, western sand cherry, or dwarf cherry. This specimen was growing near the southern margin of the Nebraaska Sandhills. Its closest neighbors were little bluestem and Scribner's panicgrass (Panicum scribnerianum= P. oligosanthes var. scribnerianum), and soapweed yucca (Yucca glauca).

Overall plant morphology (first slide) and features of entire leaders or limbs (second slide) were presented.

Lincoln County, Nebraska. Mid-June; immature-fruit stage of phenology.

25. Cherries in the hills, green ones that is- Progressively closer-in views of leaves and drupes of Bessey's or western sand cherry growing at southern margin of Nebraska Sandhills. This was on the same plant that was introduced in the immediately preceding pair of slides.

Prunus species are in the Prunoideae or Amygdaloideae subfamily of Rosaceae. Th drupes of this subfamily have structures consisting of both 1) hard, boney, or rock-like endocarp and 2) seed traditionally referred to as a stone or pit. The technical name for this stone or pit is pyrene (Smith, 1977, ps. 149, 305, 309).

Lincoln County, Nebraska. Mid-June; immature-fruit stage of phenology.

26. Another sandy (and even rosier) shrub- Prairie rose (Rosa arkansana) in central Nebraska Sandhills. In contrast to Prunus species (eg. sand cherry treated immediately above) which are in the Prunoideae (prune subfamily) and charactrized by the drupe fruit type, Rosa species are in the Rosoideae (rose subfamily) in which the fruit type is an aggregate fruit. This aggregate fruit is a form of false fruit which in Rosa species is a hip (see immediately below).

This plant was growing in moist sand.

Thomas County, Nebraska. Mid-June; plant at mid-bloom stage.

27. Roses are for lovers- Inflorescence of prairie rose in central Nebraska Sandhills. The rose ovary is within a floral cup or floral tube which is the organ formed from fusion of perianth and androecium (staminal structure) that surrounds the pistil and ultimately forms the hip (Smith, 1977, ps. 148, 298). This hip, then, is the entire fruit-bearing structure which is a form of false fruit. Within the hip is borne the actual fruit which, strictly speaking, is an achene (several achenes per hip).

Copulating insects were some species of longhorn beetle of family Cerambycidae. Any viewers who can identify this speacies please inform the author.

Technical note: score another messed up slide to Epson Perfection 700. That imprecise machine never would scan correctly the color of rose petals in second of these two slides. Color of petals was more closely copied in the first slide of these slides.

Thomas County, Nebraska. Mid-June; fully open flower.

28. Scribnage in the Sandhills- A single sprawling of Scribners panicgrass (Panicum scribnerianum= P. oligosanthes var. scribnerianum) growing on degraded tallgrass prairie at southern extremity of Nebraska Sandhills. This is one of eight species of the cool-season, C3 photosynthesis, winter rosette -forming panicgrasses, the so-called dichanthellum panicums, native to Nebraska (Kaul et al, 2006, ps. 691-695). In Hitchcock and Chase (1950, ps. ) Dichanthelium was a subgenus of Panicum. Some later authors including Gould and Shaw (1983, ps. 228-230), Great Plains Flora Association (1986, ps.1156-1163), and Barkworth et al. (2003, ps. 406-450) in the definitive Flora of North America elevated Dichanthelium to genus status. Orthers such as Diggs et al. (1999, ps. 1291-1302) and Kaul et al (2006, ps. 691-695) retained the rosette panicgrasses in Panicum.

Scribner's panicgrass is primarily an increaser on most sites in the Sandhills. It is well-adapted to heavy grazing given its relatively low stature. Scribner's panicgrass has a large biological range growing from Texas north to southern Canada from Ontario to British Columbia.

Lincoln County, Nebraska. Mid-June; early grain stages.

29 Setting grain in the Sandhills- Sexual shoots of Scribner's panicgrass produced a super crop of caryopses in the souther edge of the Nebraska Sandhills an ususually wet year. Lincoln County, Nebraska. Mid-June; early grain stages (mostly milk-stage)..

30. Sandhill spring- Nebraska Sandhills with vernal society. This range is textbook Four Horsemen (big bluestem, Indiangrass, switchgrass, and little bluestem) plus prairie sandreed tallgrass prairie in estival-autumnal aspects, but at time of this photograph it was the vernal society/aspect. The conspicuous cool-season perennial grass was Junegrass (Koleria cristata). In this vernal society Junegrass was the associate to the dominant needle-and-thread (Stipa comata).

Several of the grassland range types have some of the most pronounced seasonal societies (different seasonal plant communities) of any natural (climax) vegetation in North America. Societies (climax seasonal plant communities) of the tallgrass are more pronounced in the Nebraska Sandhills because this rangeland cover type (SRM 602; Bluestem- Prairie Sandreed) occurrs far enought north and at extreme western margin of tallgrass prairie so that there is more of a mixture of cool- and warm-season species as well as a greater array of grasses within the various subfamilies of the Gramineae. For example, botanical diversity is considerably greater in the Nebraska Sandhills than in similar tallgrass prairie vegetation of the Flint Hills and Osage Cuestas of Kansas and Oklahoma.

Logan County, Nebraska. Mid-June; late vernal aspect. FRES No. 39 (Prairie Ecosystem). K-67 (Nebraska Sand Hills Prairie). SRM 602 (Bluestem- Prairie Sandreed). Bluestem "Tall-Grass" Series, 142.11 of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Generic range site: Sands (Soil Conservation Service, 1965). Nebraska Sand Hills- Sand Hills Ecoregion, 44a (Chapman et al., 2001).

31. Sterotypic views- Textbook presentations of the Nebraska Sandhills tallgrass prairie in vernal s aspect (ie. with vernal vegetational society). Needle-and-thread was the seasonal dominant and Junegrass was seasonal associate (= dominant and associate species in vernal society). By late July through to frost the dominants would be (variously) big bluestem, switchgrass, Indiangrass, and prairie sandreed. Little bluestem was also present, but at this northward latitude little bluestem starts to become a midgrass and is clearly secondary, especially to big bluestem, sand bluestem, and prairie sandreed, the defining dominants.

A robust plant of needle-and-thread was sillhouetted against the hindquarters of lactating beef cow whose symetrical udder with a level floor and perfectly placed teats should be the envy of 90% (or more) of all college coeds, and they cow ain't fat.

32. Inside vernal verdent dunes- Interior of Nebraska Sandhills tallgrass prairie in late spring. Such grasslands are beautiful at all spatial scales. The first of these slides presented a landscape scale view while second slide showed local scale at top of a tall, grass-covered sand dune. This range vegetation was a big bluestem-sand bluestem-prairie sandreed tallgrass prairie with its vernal society dominated by such cool-season grasses as needle-and-thread and Junegrass, the two co-dominants on this range at this time.

The main spring forb was common pepperweed (Lepidium densiflorum).

The conspicuous prairie forb with white flowers in the second slide was soapweed or soapweed yucca (Yucca glauca). Pale (sun-bleached) tall shoots in foreground of second slide were switchgrass which, for whatever reasons, thrives best on top of dunes as well as the more common preferred habitat of low-lying wet areas.

33. Another two-scale spring view- Nebraska Sandhills tallgrass prairie with vernal society at combination local and landscape scales (first slide) and local scale only (second slide). On this less mesic habitat that was mostly a Sands range site (foreground) to Choppy Sands range site (distant background) little bluestem was the dominant and prairie sandreed the associate species. Junegrass was conspiuous being at peak bloom in this vernal society. In the first of these two slides the more prominent forb was Carolina puccoon or plains puccoon (Lithospermum caroliiense) in immediate foreground. The most conspicuous nongrass species in the first slide (left side of dune on right) was New Jersey tea (Ceanothus herbacea var. pubescens). The showy forb in center foreground of second slide was prairie thistle (Cirsium canescens).

34. Lush vegetation atop a sand dune- Range plant community of tallgrass prairie on one of the larger and taller dunes in Nebraska Sandhills. Little bluestem was the dominant and prairie sandreed was the associate species on the upper portion of this massive sand dune. Big and/or sand bluestem and switchgrass were other tallgrass species. Junegrass and needle-and-thread were local associate species, at least in this vernal aspect/society.

The prominent and dominant broadleaf species was New Jersey tea. Prairie thistle was present as a showy though limited forb.Other forbs included Akansas rose (Rosa arkansana) and soapweed or soapweed yucca, two species interpreted as subshrubs by some workers.

35. To each its own season- Needle-and-thread was the dominant of the vernal society of a tallgrass prairie on a Choppy Sands range site in the heart of Nebraska Sandhills. An early morning sky provided the backdrop for this species that is only an associate to the overall dominants of this range type which included little bluestem (the local dominant), sand and/or big bluestem, and prairie sandreed. Prairie Junegrass was another important member of the vernal society (locally the cool-season associate to needle-and-thread). Blue grama was an important shortgrass component of this unquestionably tallgrass prairie.

36. Our turn- The sward of a needle-and-thread consociation in the vernal society of Nebraska Sandhills tallgrass prairie. Societies (different combinations of plant species that vary by season within a given climax vegetation; seasonally variable climax plant communities) are often very pronounced in grasslands. This is more the case in mid-latitudes where winters are colder while summer temperatures are similar (though of shorter duration) to those far to the south. The result is that there are cool-season and warm-season dominants in quite pronounced societies. For comparison, this is more the case for tallgrass prairie in the Sandhills of Nebraska than in the Flint Hills and Osage Cuestas of Kansas, Oklahoma, and Missouri.

As the current author applied society in this description it referred to the seasonal aspect rather than that of vegetational layers (Clements, 1936; Weaver and Clements, 1938, ps.96-99).

The estival (summer) society of this tallgrass prairie on a Choppy Sands range site was dominated by little bluestem with sand and/or big bluestem and prairie sandreed as associate species. Peak standing crop occurs during the estival society. Ultimately, needle-and-thread was eclipsed by the warm-season tallgrass species. For now, however, in the morning beauty of late spring needle-and-thread was in center stage and sharing the limelight with none of the summer-shining phytofolk.

37. June in the Nebraska Sandills- Junegrass or prairie Junegrass (Koeleria cristata= K. pyramidata= K. macrantha= K. nitida) in late spring growing at two locations on a Choppy Sands range site in tallgrass prairie of the Nebraska Sandhills. These two specimens were at peak standing crop and full bloom (anthesis) showing their species to "maximum effect".

Junegrass is one of the most widely distributed prairie grasses in North America being a member of tallgrass prairie, true prairie, mixed prairie, Palouse Prairie and Pacific prairie as well as pinyon pine-juniper woodland and ponderosa pine forest range. For example, Junegrass has been found in almost all counties of Nebraska (Kaul et al, 2006, p. 679) and doubtless is native statewide. With such a vast biological range there is considerable morphological --not to mention genetic-- variation in this species. This is especially the case for its panicle inflorescence (see immediately below). Barkworth et al. (2004, p. 754) noted that Junegrass is "a polymorphic, polyploid complex". 'nough said.

Junegrass is one of the "big 200" range plant species on the International Range Plant Identification Contest sponsored by the Society for Range Management (Stubbendieck et al., 1992, ps. 64-65), proof of the importance of this grass on North American ranges. The Range plant Handbook (Forest Service, 1940, p. G76) is a standard reference for Junegrass. The field guide of Skinner (2010) is outstanding for identification, description, and superb photographs of Junegrass: highly recommended.

Kaul et al. (2006, p. 679) stated that the older speccific epithet cristata was "illigitimate". So what? What makes it "illigitimate" when cristata was used in the early classic works such as manuals or floras, including Hitchcock and Chase (1950), the Range Plant Handbook (Forest Service, 1940), and Fernald (1950, 145), as well as in the seminal ecological treatments like Weaver (1954) and Weaver and Albertson (1956)? How can it be "illigitimate" when it when it was on the birth and baptisim certificates (at least the copies available to rangemen)? Barkworth et al. (2004, p. 754) asserted that macratha was the proper specific epithet for North American Junegrass whereas K. pyramidata was restricted to Eurasia. Of course K. pyramidata was the previous correct or proper species designation. Nobody short of God knows what will replace the epithet macratha, but it is a safe bet that some mothball brain-vapored taaxonomist will assert that such and such is right and so and so is "illigitimate". Ought to be good for another publication don't you think.

Thomas County, Nebraska (first slide) and Logan County, Nebraska (second slide). Mid-June; peak-bloom phenological stage.

38. Junegrass panicles- Although Kaul et al. (2006, p. 679) described Junegrass in Nebraska as having a "congested panicle", which has been the standard description including McGregor et al. (1986, p. 186) this is not always the case even within this one state let alone the general Great Plains Region. As was the case more often than not, Hitchcock and Chase (1950, p. 281) more accurately descried the Junegrass panicle as "...spikelike, dense (loose in anthesis), often lobed, interrupted, or sometimes branched below...". The features of lower branching, lobed and loose or more open at flowering were all clearly presented in these examples.

Loup County, Nebraska. Late June; immediate post-anthesis.

Location note: other photograaphs and descriptions of Junegrass are scattered variously throughout this publication, including Tallgrass Prairie (Interior)-IB and Juniper-Pinyon Woodland.

39. Southern extremity of Nebraska Sandhills- Landscape of climax tallgrass prairie in Sandhills of Nebraska on southern edge of this range type where it joins the Loess Hills mixed prairie type. Little bluestem was the dominant on tops and upper sides of ridges while switchgrass dominated the bottoms of ridges and small basins situated among ridges. Shoots of switchgrass were most prominent in foreground of the second of these three slides and, to lesser degree, in foreground of first slide. Big and sand bluestems and prairie sandreed were associate tallgrass species. Needle-and-thread was an associate in this vernal society, but it was not a dominant at ultimate expression or peak standing crop in estival/autumnal societies and aspects.There were some short grass species, especially blue grama with lesser amounts of buffalograss. Sixweeks fescue (Festuca octoflora), a native "sixweeks grass" (ephemeral), was the only important annual species. Japanese chess (Bromus japonicus), a naturalized Eurasian annual grass, was present in trace amounts.

Midgrass species were not abundant enough to form a continuous layer in this range plant community thereby forming a mixed prairie. This was undoubtedly tallgrass prairie though it was postclimax (Weaver and Clements, 1938, ps. 86, 102, 462, 520-521) in this semiarid precipitation zone. Remarkable water-holding capacity of the sandy soil was the determining factor that permitted development of this postclimax range vegetation that would otherwise have been mixed prairie..

Prairie forbs included prairie pussytoes (Antennaria parvifolia), prairie larkspur (Delphinum virescens), and plains or Carolina puccoon.

This tallgrass prairie was in Excellent condition class. It was presented as another example of the ultimate development and expression of the potential natural vegetation of Nebraska Sandhills grassland.

Lincoln County, Nebraska. Mid-June; late vernal aspect. FRES No. 39 (Prairie Ecosystem). K-67 (Nebraska Sand Hills Prairie). SRM 602 (Bluestem- Prairie Sandreed). Bluestem "Tall-Grass" Series, 142.11 of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Generic range site: Sands (Soil Conservation Service, 1978). Nebraska Sand Hills- Sand Hills Ecoregion, 44a (Chapman et al., 2001).

40. Six weeks in the Sandhills- Sixweeks fescue (Festuca octoflora= Vulpia octoflora) growing on the southern Nebraska Sandhills range described immediately above. This is a native annual grass, one of the very few of North American grasslands. Sixweeks fescue is a widely distributed species with a biological range that extends across Canada from Atlantic to Pacific and into Mexico and from the Sonoran (Colorado) Desert of California across down to Florida.

Following Fernald (1950, p. 108) some flora or manuals placed the annual fescue species in genus, Vulpia. This view was followed in the Nebraska flora (Kaul et al, 2006, p. 720). Other flora, including the definitive source for the general Great Plains region (Greaat Plains Flora Association, 1986, p. 1180) retained the annuals in the traditional Festuca.At time of this writing there did not appear to be a good source that treated sixweeks fescue from other than a taxonomic view other than, of course, Pasture & Range Plants (Phillips Petroleum Company, 1963, p. 42; Fort Hays State University, 2006, p. 42) which incidentially from first to second edition changed from Festuca to Vulpia. Barkworth et al. (2006, ps. 448-454), the consummate splitters, also used Vulpia for the annual fescues as was 100% predictable.

Hylton and Bement (1961) described growth and development of sixweeks fescue from germination/emergence to grain-ripe. They found that grazing, especially heavier degrees of defoliation, benefitted growth and populations fo this annual grass. Seeds of sixweeks fescue germinated and seedlings emerged in autumn whereupon plants went into dormancy and resumed growth in late winter or early spring (Hylton and Bement, 1961) .

This annual does not furnish--at least, as a general rule--the quality or quantity of forage as that from perennial grasses. (Hyder and Bement, 1963) stated that on the Central Great Plains "[s]ixweeks fescue is unacceptable to cattle at all seasons". There are undoubtedly exceptions under some range situations such as perhaps unusually moist conditions and resultant lush herbage or rapid recovery from drought at which times any range feed can prove priceless. Also, sixweeks fescue often provides vegetative soil cover which also can be priceless in sandy land like the Nebraskas Sandhills. On the other hand, this ephemeral grass has been described as interfering with utilization of more nutritious forage species such as blue grama because cattle avoid range areas heavily populated by sixweeks fescue (Hyder and Bement, 1963).

Lincoln County, Nebraska. Mid-June; mature and senescing (dying) phenological stages with soft to hard dough state of caryopses.

41. Breeze-blown and ripening- Wind-blown image of panicle with spikelets of sixweeks fescue growing on a range at southern extent of Nebraska Sandhills. Number of florets per spikelets in this species varies from seven to eleven or, sometimes, thirteen (Great Plains Flora Association, 1986, p. 1180). Wind is the bane of the range plant photographer, and in this case somewhat wind-blown images could not be avoided due to need for slow shutter speed for depth of field details combined with a stiff sandhills breeze. Ironically, the wind-blown effect did not show anemochory, dispersal of plant propagules or germules by wind, because this species is not cross-polinated. Sixweeks fescue, like the other annual Festuca (Vulpia) species, is cleistogamous (Lonard and Gould, 1974), the anthers are not exerted and instead individual flowers (florets) are self-fertilized. Interesting little range plant.

Lincoln County, Nebraska. Mid-June; mature and senescing (dying) phenological stages with soft to hard dough state of caryopses.

42. Plains or Carolina puccoon (Lithospermum caroliniense)- Plains puccoon is one of the earliest blooming forbs over much of the Nebraska Sandhills. This far north that earlier blooming was late spring (in fact almost early summer). Plains puccoon is also one of the most widely distrubuted forbs in Nebraska. The specimen presented in these three successively closer-in views was growing on the Nebraska Sandhills tallgrass prairie range introduced in the preceding three-slide set.

Lincoln County, Nebraska. Mid-June; obviously full-bloom phenological stage.

43. NOut on a lark- Prairie larkspur (Delphinum virescens, often included with D. carolinianum [see next succeeding caption]) growing on southern extreme of Nebraska Sandhills. It was remarked above that this was an important forb along with Carolina or plains puccoon and prairie larkspur on example rangeland presented here. The critical thing about prairie larkspur from perspective of range and pasture management is its toxic property. Readers were referred to the two standard references for details of larkspur poisoning: Kingsbury (1964, ps. 131-140 passim) and Burrows and Tryl (2001, ps. 1009-1017 passim).

This sole plant of its species in this immediate area was growing in barrow ditch outside a cattle pasture of climax tallgrass prairie (Excellent range condition class) that was shown and desribed shortly above. At the abuncance of prairie larkspur on this grassland livestock poisoning would be nearly impossible (even if this particular plant had been inside the pasture).

The Delphinum species have traditionally been categorized based on their mature heights and placed into three groups: 1) low larkspurs, 2) intermediate larkspurs, and 3) tall larkspurs (see again above references). D. virescens is one of the intermediate larkspurs.

Lincoln County, Nebraska. Mid-June; peak standing crop and full-bloom phenological stage.

44.NLike on a lark- A few flowers on the inflorescence of plains larkspur in the southern Nebraska Sandhills. These flowers were on the same plant tht was shown in the immediately preceding slide. D. virescens is very similar to D. carolinianum from which it can be distinguished by various features such as those of pedicels (McGregor et al., 1986, p. 94). Features between these two taxa are so similar that some authorities proposed that D. virescens be interpreted as a subspecies of D. carolinianum; however, D. virescens forms a complex with several other species and their taxonomy warrents further sudy (McGregor et al., 1986, p. 94).

The common name "larkspur" was derived from the imagined similarity between the backward pointing toe (spur) of a lark and the elongated receptacle of Delphinum species on which corolla and calyx are inserted. The perianth (calyx and corolla taken together) is typically petaloid (resembling petals) and not differentiated into these two floral units (Smith, 1977, p. 80). This is a consistent feature of the Ranunculaceae, buttercup family.

Lincoln County, Nebraska. Mid-June; full-bloom phenological stage.

45. Girls' night (and day) out- A plant of prairie, small-leaf or little-leaf pussytoes plant in full flower. This plant was growing on a wheatgrass-needlegrass true prairie range in the sedimentary unglaciated Northern Great Plains. This flowering plant of Antennaria parvifolia was included here to complement another plant of littleleaf pussytoes that was at fruit-shatter stage growing on this sandhills tallgrass prairie.

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

48. Thriving in the sandhills- Single plant of prairie or Platte thistle (Cirsium canescens) in the heart of the Nebraska Sandhills. Associated range plants were sand bluestem and prairie sandreed. This native composite is not a weed--at least under most situations--on properly managed tallgrass prairie in the Sandhills parardise. Platte thistle is found on virgin grasslands of the general Sandhills Region as an infrequent species with only one to a few individuals at very wide intervals (ie. very low density).

In spite of its bone-colored flowers (versus bright pink or red of other Cirsium species) Platte or prairie thistle is a beautiful plant. The specific epithet, canescens, means "grey-pubescent" which could apply to the stem (and some leaf surfaces) as well as the more conspicuous head. Even this duller color of flower could only subdue and not negate the showiness, prominence, and attractiveness of this native grassland forb.

This particular (and quite robust) individual had grown to large size (it had a height of 30 inches, which could serve as a guage to breadth of the plant) in an extremely wet spring that followed an equally wet winter.

Cherry County, Nebraska. Mid-June; peak bloom stage.

49. Platted in the sandhills- A plant of Platte or prairie thistle (first slide) with closeup of two of its heads (second slide) in Nebraska Sandhills. This specimen was growing amid prairie sandreed and sand bluestem. The first slide presented three heads, two of which were still at peak bloom and one that was "spent" (had flowered) at immediate post-bloom stage. The second slide showed one head at peak bloom (left) and one head at immediate post-bloom (right) stage. These two blooms were the two heads at left in first slide.

The more botanically precise term for head is capitulum (plural, capitula) which is the inflorescence type of the Compositae. In capitula numerous to many individual flowers or florets are inserted on a common receptacle, known as a disk, from the perimeter of which project phyllaries (bracts) that resemble sepals of other flower types (Diggs et al., 1999, p. 287). The entire capitulum resenbles a single large flower, though in reality it consist of dozens of small flowers (the florets).

Characteristic leaf shape was presented in the first slide.

Logan County, Nebraska. Mid-June; peak bloom stage.

50. Heads up in the sandhills- Two capitula (heads) of Platte or prairie thistle in the Nebraska Sandhills. The Compositae capitulum was described in the preceding caption. The vase- or goblet-shaped base of the receptacle, bears spines the bases of which somewhat resemble shingles on a roof or plates of armour. The receptacle is the "expanded apex of the pedicel upon which the flower series are inserted" (Smith, 1977, p. 306).

The adjective and specific epithet, canescens, means "grey-pubescent" which could refer to the pubescence of the stem (as shown in the second slide) as well as the bone-colored capitulum. Even allowing for the weediness or noxious feature of most thistles they are remarkably attractive plants. Though sporting a comparatively dull capitula (compared to the bright reds, pinks, lavenders of other thistles), Platte or prairie thistle was still an attractive plant--even by the high standards of many range forbs. Furthermore (and as explained above) this native thistle is no weed, at least not under conditions of proper range management.

Logan and Thomas (first and second slide, respectively) County, Nebraska. Mid-June; peak bloom stage.

51. Big and bold in the Sandhills- Big or large beardtongue (Penstemon grandiflorus) growing in central Nebraska Sandhills on a tallgrass prairie range dominated by sand bluestem, prairie sandreed, and little bluestem. As indicated by the common name this is one of the largest of the many Penstemon species. Some of the shoots in these photographs were over a yard in height, and much of that height had the bright lavendar flowers blooming along them. .

Good references for the Penstemon species on central and northern portions of the Western Range was Rickett (1973, p. 592-609), but limited coverage and even of photographs (there was not a color plate of P. grandiflorus).

Thomas County, Nebraska. Late June; peak standing crop, peak-bloom stage.

52. Big, bold blossoms- Sequential views of inflorescence and individual flowers of big beardtongue in Nebraska Sandhills. The petals of this Penstemon species are some of the showier and brighter of all the beardtongues, of which there are numerous species on the Western Range. The specific epithet grandiflorus was obviusly most apropriate for this prairie forb.

Thomas County, Nebraska. Late June; peak-bloom stage (and no question about that).

53. Climax sward of Sandhills tallgrass prairie- Turf of vernal society of climax Nebraska Sandhills tallgrass prairie. Warm-season (and overall) dominant species at local scale was switchgrass (the conspicuous taller and bleached-out shoots) with prairie sandreed, big and/or sand bluestem, and little bluestem associate species. Needle-and-thread, the dominant cool-season species, was at peak standing crop with grain in milk to soft-dough stages. Buffalograss was preparing to bloom. Cover, relative density, and general abundance of this midgrass and shortgrass species were "not enough" to classify this grassland as mixed prairie.

This was textbook postclimax tallgrass prairie in the semiarid precipitation zone of westcentral Nebraska (Great Plains physiographic province). It was mesic grassland that developed due to greater water-holding capacity of sandy soil; hence, it was postclimax (ie. regional edaphic conditions "overrode" the usual dominant role of climate. This was postclimx versus climax (= climatic climax) as viewed from perspective of monoclimax theory (Clements, 1916; 1936). From the polyclimax theory, often attributed to Arthur Tansley, such tallgrass prairie vegetation in the Nebraska Sandhills would be an edaphic climax.."Same difference."

54. Little "sand dune" due to a little burrower- Mound of soil from the burrow of Great Plains pocket gopher (Geomys bursarius). Great Plains pocket gopher is a widely fossorial (literally "burrowing"; hence, living underground and generally having such adaptations as limbs and feet adapted to digging) mammal with a species range extending from the Canadian Prairie Provinces south into Mexico (Case and Jasch, 1994). This rodent is not only widely distributed it, also thrives in a variety of grass-dominated plant communities ranging from climax grasslands to golf courses, graveyards, and front yards. Diets of Great Plains pocket gophers are overwhelmingly of herbaceous plants,including the three categories of grasses, grasslike plants, and forbs (Luce et al, 1980).

This particular mound was on the range of climax Nebraska Sandhills tallgrass prairie that was in Excellent range condition class described in the two immediately preceding slide-caption sets. This particular mound and associated burrowing was in the gateway into this pasture where vehicle disturbance had eliminated most of the perennial plants so as to be almost exclusively sixweeks fescue, a native annual (ephemeral) grass species. Probably, easier digging for these little beggars.)

Pedoturbation is the term applied to "mixing within a soil or sediment profile by various processes, such as animal borrowing , tree throw, freeze-thaw cycles; [i]t usually involves disturbance of the skeletal fabric as opposed to redistribution of only the fine particles (Soil Science Society of American, 2001). This was thus an example of organisms as a soil formation factor.

Lincoln County, Nebraska. Mid-June

55. Mixed prairie grazing disclimax- Overuse over course of several summer and early autumn seasons by beef cattle had converted a bluestem-prairie sandreed Nebraska Sandhills tallgrass prairie into a mixed prairie of cool-season midgrass species (both decreasers, like needle-and-thread and Junegrass, and increasers, notable a rosette panicgrass) and warm-season shortgrass species (pirmarily blue grama with some buffalograss). The rosette Panicum species was Scribner's panicgrass (P. scribnerianum).

Long-term overuse (excessive defoliation; grazing harvest that exceeds sustained yield) is overgrazing. This former tallgrass prairie range had obviously been overgrazed (was substantially overstocked for numerous grazing seasons) and converted into a range of midgrass species. Such an anthropogenic (human-caused) type conversion was a grazing disturbance climax, a man-made range plant community that would persist so long into the future that it resembled the dynamic equilibrium of an actual climax (Weaver and Clements, 1938, ps.86-88).

Contrast this range vegetation to that on the same (or very similar combinations of range sites) presented in preceding slides.

Students should note the numerous mounds of soil (foreground in first slide) from burrowing by Great Plains pocket gopher. (See immediately preceding photograph for close-up of this form of pedoturbation and detail of burrowed soil heap).

56. Depleted Sandhills sward- Tallgrass prairie (formerly) at southern margin of Nebraska Sandhills degraded by overgrazing during the warm-growing season had resulted in almost complete loss of climax warm-season tallgrass species like big and sand bluestem, switchgrass, and prairie sandreed. In a textbook example of range deterioration cool-season climax dominants like needle-and-thread and Junegrass remained during the spring and early summer seasons along with warm-season associate species such as blue grama whereas the warm-season (and overall range type) dominants were grazed out.

A striking example of this replacement of warm-season species by cool-season grasses was loss of switchgrass concomittantly with increase in the increaser panicoid grass, Scribner's panicgrass (Panicum scribnerianum). Whereas switchgrass is a C4 species (one that ultimately fixes photosynthetate as a four-carbon intermediary) Scribner's panicgrass is a C3 species that exclusively utilizes the Calvin-Bensen photosynthetic pathway to fix photosynthate in threee-carbon units.Some agrostologists such as Gould (1975) and Barkworth et al. (2003) transferred these cool-season, C3 panicgrasses that form winter rosettes from the Panicum genus to of Dichanthelium which was a subgenus of Panicum as treated by Hitchcock and Chase (1951). One plant of Scribner's panicgrass was displayed in immediate center foreground of this slide.

The sward of this grazing disclimax can be compared to the sward (presented three slide-caption sets above) of climax vegetation on a cattle range in Excellent condition class that had essentially the same range site(s) as the range site and precipitation zone of this disturbance climax.

57. Wonder of the Nebraska Sandhills- Sand dune tallgrass prairie dominated by prairie sandreed and big and/or sand bluestem, little bluestem, and switchgrass (fore back to near background) with a large freshwater lake and grasslike species (presented below) gleaming on the horizon (distant background). Some of this wetland range vegetation was also in midground where a local depression was dominated by common threesquare or chairmakers' rush (Scirpus pungens).

It is the marvelous diversity in grassland and marsh communities--often within remarkably short distances of each other--and the occurrence of surface water in the semiarid zone that makes the Nebraska Sandhills such a "wonderment". Add in the grazing capacity and raw beauty of miles of virgin grassland and the Sandhills become a "pure dee wonderment".

These two landscape-scale views presented the physiogonomy of the postclimax (= edaphic climax) tallgrass prairie at their zenith in the Sandhills.

58. Pristine sward- Perspectives of physiogonomy at close range along with structure and composition of Nebraska Sandhills tallgrass prairie. This range vegetation was a "closer-to-it" look at the range plant community introduced in the immediately preceding photographs. Plant cover was alomst exclusively that of the tallgrass species of prairie sandreed, big and/or sand bluestem, switchgrass, and little bluestem.

This climax grassland community was on top of a ridge (narrow top or edge of a low sand dune; see again preceding two slides) so that there were no aquatic, woody, or grasslike plants present.

59. Compostiion and structure of the pristine sward- Two "photoquadrants" of the climax tallgrass prairie vegetation presented in the two preceding two slide-caption sets. These were the "closest-in" looks at this range plant community that permitted simultaneous observation of plant species and their relative cover and density, general abundance, etc. along with layering and spatial arrangement of the sward.

The taller, tan or buff-colored, cespitose plants were the bluestems. There were also a few whitish or bleached-out shoots of switchgrass. The lowest (closest to the ground) shoots with broad, glaucous-like leaves were prairie sandreed. The second of these two slides was a close-up of prairie sandreed plants shown at greater distance at right in the first slide.

60. Atop a grassy dune- Landscape-scale look across Nebraska Sandhills tallgrass prairie with a large (by standards of a natural body of water in the semiarid zone) freshwater lake in the background. Grassland vegetation was the same as shown in previous slides: prairie sandreed, big and/or sand bluestem, little bluestem, and switchgrass.

Wetland (marsh) vegetation of the lake and overflow or drainages into it were featured in the next five (5) slide-caption sets.

61. Where dry and wet meet- Dry--less mesic--upslope or dune top with tallgrass prairie (foreground of both slides) going down to wet prairie (foreground of first slide and midground of second slide) and, finally, to marsh with water-covered land surface (background of first slide and midground of second slide). The small channel of water was a drainage or overflow from the large freshwater lake (in background of both slides).

Tallgrass prairie was the same that was treated in preceding slides and captions. This two-slide set along with subsequent sets treated the wet prairie and marsh vegetation of this Nebraska Sandhills landscape. Much of the herbage growing in standing water shown in second slide was bluejoint reedgrass (Calamagrostis canadensis).

Rock County, Nebraska. Late June; early estival aspect. FRES No. 39 (Prairie Ecosystem). K-67 (Nebraska Sand Hills Prairie). SRM 602 (Bluestem- Prairie Sandreed). Bluestem "Tall-Grass" Series, 142.11 of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Wetland vegetation was (would be if shown, as it was not) Bulrush or Tule Series, 242.33 of Plains Interior Marwshland 242.4 of Cold Temperate Marshland 242 of Brown et al. (1998, p. 45). Nebraska Sand Hills- Sand Hills Ecoregion, 44a (Chapman et al., 2001).

62. Lake's edge- Along perimeter of a freshwater lake in Nebraska Sandhills wet prairie dominated by switchgrass (bottom ecotype) developed gradually into marsh with progress along an environmental hydrologic gradient into the lake. Plant species of the marsh included bluejoint reedgrass, common spikerush (Eleocharis palustris), meadow willow (Salix petiolaris), and common threesquare or chairmakers' rush. Outward from the switchgrass-dominated wet prairie was margin of less mesic tallgrass prairie (immediate foreground) of prairie sandreed, big and/or sand bluestem, and little bluestem (see again above).

Rock County, Nebraska. Late June; early estival aspect. FRES No. 39 (Prairie Ecosystem). K-67 (Nebraska Sand Hills Prairie). Prairie was a variant of SRM 602 (Bluestem- Prairie Sandreed). Bluestem "Tall-Grass" Series, 142.11 of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Wetland range was Wetland vegetation was (would be if shown, as it was not) Bulrush or Tule Series, 242.33 of Plains Interior Marshland 242.4 of Cold Temperate Marshland 242 of Brown et al. (1998, p. 45).Nebraska Sand Hills- Sand Hills Ecoregion, 44a (Chapman et al., 2001).

63. Meet ya at the lake- Perimeter of tallgrass prairie and marsh at a natural freshwater lake in Nebraska Sandhills. Outermost (farthest from lake) was tallgrass prairie dominated by prairie sandreed, big and.or sand bluestem, and little bluestem. Inward (closer to the lake edge) bottomland swithcgrass formed a nearly exclusive zone of wet prairie vegetation. Inward from the swithcgrass zone was a lake's edge marsh of chairmaker's rush or common threesquare and broadleaf cattail (Typha latifolia) with a narrow zone of bluejoint reedgrass.

All these zones of range vegetation were shown in the first of these two slides. The second photograph featured the threesquare-broadleaf cattail marsh and the narrow zone of bluejoint in immediate foreground. Detailed view of the threesquare-broadleaf cattail zone and bluejoint zone was subject of the next slide...

Rock County, Nebraska. Late June; early estival aspect. FRES No. 39 (Prairie Ecosystem). K-67 (Nebraska Sand Hills Prairie). Prairie was a variant of SRM 602 (Bluestem- Prairie Sandreed). Bluestem "Tall-Grass" Series, 142.11 of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Wetland vegetation was (would be if shown, as it was not) Bulrush or Tule Series, 242.33 of Plains Interior Marshland 242.4 of Cold Temperate Marshland 242 of Brown et al. (1998, p. 45). Nebraska Sand Hills- Sand Hills Ecoregion, 44a (Chapman et al., 2001).

64. Sandhills pothole- Local depression in the varied landscape of Nebraska Sandhills was filled with water (ie. a localized marsh or prairie pothole) and supported a zone of bluejoint reedgrass (immediate foreground) with a vegetational zone immedaiately behind that was comprised almost exclusively of common threesquare or chairmakers' rush.

Rock County, Nebraska. Late June; early estival aspect. Wetland vegetation was (would be if shown, as it was not) Bulrush or Tule Series, 242.33 of Plains Interior Marwshland 242.4 of Cold Temperate Marshland 242 of Brown et al. (1998, p. 45). Nebraska Sand Hills- Sand Hills Ecoregion, 44a (Chapman et al., 2001).

65. Sloshing away from the pothole- Wet prairie (foreground of both slides) exterior to the Nebraska Sandhills prairie pothole featured in the immediate preceding photograph. The wetland vegetation of that pothole was visible in the far midground of both of these slides. Upland (dry or less mesic) tallgrass prairie range of prairie sandreed, big and/or sand bluestem, and little bluestem was barely visible in distant background of these two slides.

Wet prairie range featured in these two slides (foregrounds) was dominated by a bottomland ecotype of switchgrass with wild licorice (Glycyrrhiza lepidota) as the associate plant species. There were also a few small plants of meadow willow. Immediately behind the switchgrass-dominated wet prairie there was an even wetter habitat of wet prairie that was populated by bluejoint reedgrass and common spikerush. Behind the bluejoint-spikerush zone was the pothole of common threesquare or chairmakers' rush (far midground) that was the subject of the preceding slide.

Rock County, Nebraska. Late June; early estival aspect. FRES No. 39 (Prairie Ecosystem). K-67 (Nebraska Sand Hills Prairie). Prairie was a variant of SRM 602 (Bluestem- Prairie Sandreed). Bluestem "Tall-Grass" Series, 142.11 of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Wetland vegetation was (would be if shown, as it was not) Bulrush or Tule Series, 242.33 of Plains Interior Marshland 242.4 of Cold Temperate Marshland 242 of Brown et al. (1998, p. 45). Nebraska Sand Hills- Sand Hills Ecoregion, 44a (Chapman et al., 2001).

66. Wet prairie community up close- Two closer-in "photoplots" presented botanical composition and structure of the wet prairie featured immediately above. The first "photoplot" showed wet prairie range vegetation that was dominated by switchgrass and bludjoint reedgrass with wild licorice and meadow willow. Second "photoquadrant" was a local stand of common spikerush (Eleocharis palustris) with both wild licorice and meadow willow.

67. Back on drier prairie (with lone forbs amid tallgrass)- Plains or Carolina puccoon (Lithospermum caroliiense) shared tallgrass prairie with dominant tallgrass species of big and/or sand bluestem, prairie sandreed, switchgrass, and little bluestem. This range vegetation was outward from (away from water-side) from the wet prairie and marsh plant communities treated immediately above and all the way back to the sand dune prairie featured in the opening of this discussion.

68. Armegeddon in Nebraska Sandhills- A mere half mile from the lovely pristine Nebraska Sandhills landscape of upland prairie, wet prairie, pothole marsh, and freshwater lake there appeared this apocalyptic scene. Complete, total, absolute destruction of the Nebraska Sandhill climax vegetation by conversion to a field of dent corn (Zea mays indentata) irrigated by a center pivot system. If readers can bear to view it they will sadly note what remains of a small lake that just "yesterday" in successional time was a fabulous complex of tallgrass prairie and marsh.

In a heartbreaking play on words it weas revealed that the Four Horsemen of the Prairies with prairie sandreed as a fifth range rider had been analiated and replaced by a domestic species that most agrostologist treat as a member of the Andropogoneae, the bluestem tribe.

Sadest of all--if viewers can force another glance--is that the highly erodible soil of the Sandhills was left vulnerable to the forces of erosion by the sinful specter of clean tillage which should have been abandoned decades ago.

Rock County, Nebraska. Late June. The Apocalypse.

69. Words could not suffice- There were not words powerful enough to convey the destruction of the virgin tallgrass prairie of the Nebraska Sandhills for production of dent corn on obviously marginal and highly erodible land. Center pivot irrigation combined with clean tillage (ie. no stubble or trash left on the soil surface for protection against erosion) had replaced the virgin sod of tallgrass prairie and marshes while a mud hole was all tht was left of a small freshwater lake.

This sad apocalyptic sight was only a half mile from the pristine Sandhills landscape featured above. The X-rated pictures (for nudity of the land and profane treatment of God's creation) was begrudgingly presented in the interest of truthfulness as to what is happening to far too much of the treasured Sandhills. Urban sprawl is yet another reason for loss of too much of "God's own cow country".

Perhaps this owner or renter sought to capitalize on the boondoggle and blantantly pork barrel, buy-votes scandal of corn subsidy for ethanol production, one of the most horrid, ill-advised projects yet to come out of the United States Congrsss. And they knew better; do not even think twice otherwise.

Rock County, Nebraska. Late June. Armegeddon.

70. Or was that Armegeddon?- A former cornfield planted back to big bluestem, sand bluestem, litttle bluestem, Indiangrass, and switchgrass under the Conservation Reserve Program. Man, the Manipulator of Ecosystems has the power to create or destroy, to build or to demolish, to conserve or to waste.

Lincoln County, Nebraska. Mid-June.

71. Not the final fight, but a victory field for now- Details of a Conservation Reserve Program range reseeding project. Outstandingly successful seeding project in Nebraska Sandhills that resulted in establishment of big bluestem, sand bluestem, little bluestem, Indiangrass, and switchgrass on what a two years before had been a field of dent corn. Note remnants of cobs and stalks in both of these slides.

Lincoln County, Nebraska. Mid-June.

72. Victory at Armegeddon- From the stubble of a former cornfield plants of big bluestem, sand bluestem, little bluestem, Indiangrass, and switchgrass grew out of a grassland restoration project in Nebraska Sandhills. A range reseeding project under auspices of Conservation Reserve Program established tallgrass species on a former corn field with the outstanding results seen here.

The rewarding outcome of grass establishment from this reseeding was obvious at end of second spring following planting. Corn cobs and stalks from the last crop of dent corn that was harvested two autumns ago added a "touch of class" and laughed at the irony portrayed by this project and, for that matter, the entire Conservation Reserve Program.The ironic turn of events and paradoxical twist seen here is that landowners who were the worst stewards of what was entrusted to them were the very one to benefit from a government handout that was doled out in the name of Conservation. Jesus' "good and faithful servant" reaped less reward for his faithfulness than the faithless servant who digged in the ground and buried what was entrusted to him. A wicked and cruel finale to the saga of the sodbuster and society.

Fair? No. Just? No. Any alternative once the prairie land was "broke out'? No, not that this rangeman could see. Would it more more fair, just, equitable, and of greater benefit to society if abandoned marginal land was left to erode away in order to punish the unfaithful servant? No, it would only compound the market failure; only expand the negative externality so as to extend greater harm to more people over a longer period of time. Hobson's choice in hard times. Payment for retirement and reseeding of highly erodible land was the lesser of two evils.

Either way, the prairie tallgrass species had the last laught--at least for now.

Lincoln County, Nebraska. Mid-June. Another chance.

73. Does not have to be spectacular to be dramatic- Under an overcast sky ther rolled this landscape scene of Nebraska Sandhills with shallow depressions or basins situated among slightly higher sand dunes. In this view, with cloudiness of a passing early morning shower, a basin (seen as a narrow strip of green in the midground) lay between the lighter green and tan combination of a bluestem-prairie sandreed tallgrass prairie. Topography of the Sandhills is not of breathtaking grandeur like some immense canyon or towering mountain peaks yet changes in natural vegetation due to microrelief are frrequently profound. Details of this tallgrass vegetation were presented in the next slide.

74. Where it is drier- In the Nebraska Sandhills at top of low sand dunes that rose (ever so slightly) above long, narrow basins or depressions there was a climax tallgrass prairie of prairie sandreed, big and/or sand bluestem, and little bluestem. A short early morning shower was in the offing which offered the promish of precious water and gave a different aura to the climax grassland community of this Excellent condition class range.

75. Standing water yet not a lake- A narrow basin or depression in the Nebraska Sandhills filled with water that percolated from higher sand dunes and then this infiltration water drained into the lower basins by subsurface flow. There was likely some surface runoff (overland flow) from rainfall and, perhaps, snowmelt, but much of the water in this basin resulted from subsurface latral flow. Hydrology of the Nebraska Sandhills is at the heart of their unique tallgrass prairie and marsh plant communities within the semiarid zone.

The top and upper sides of sand dunes were dry or upland tallgrass prairie dominated by little bluestem, sand bluestem, and prairie sandreed (foreground of first slide; distant background of both slides). The basin or depression and bottom of dunes was wet prairie dominated overall by bluejoint reedgrass, but with switchgrass and common spikerush locally dominant (otherwise associates to bluejoint).

This was ephemeral wet prairie with local or microrelief being such that this narrow band of wet habitat dried out by mid to late summer depending on storage of soil water and/or quantity of summer rainfall.

Rock County, Nebraska. Late June; early estival aspect. FRES No. 39 (Prairie Ecosystem). K-67 (Nebraska Sand Hills Prairie). Prairie was a variant of SRM 602 (Bluestem- Prairie Sandreed). Bluestem "Tall-Grass" Series, 142.11 of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Wetland vegetation was (would be if shown, as it was not) Bulrush or Tule Series, 242.33 of Plains Interior Marshland 242.4 of Cold Temperate Marshland 242 of Brown et al. (1998, p. 45). Nebraska Sand Hills- Sand Hills Ecoregion, 44a (Chapman et al., 2001).

76. Beauty of and on Nebraska Sandhills prairie- A striking herd of Hereford replacement heifers (and one very lucky bull) grazed a low sand dune covered by climax tallgrass prairie in which little bluestem and an upland ecotype of switchgrass with prairie sandreed the associate.

The heifers had been dehorned, but horns of the bull had only been tipped. The fancier of horned Herefords who photographed this scene wondered why the "mutilitation" as these were undoubtedly registered or, at least, purebred Herefords.

Certainly there is nothing more beautiful than quality livestock on climax grassland in Excellent range condition. Enjoy.

Loup County, Nebraska. Late June; early estival aspect. FRES No. 39 (Prairie Ecosystem). K-67 (Nebraska Sand Hills Prairie). SRM 602 (Bluestem- Prairie Sandreed). Bluestem "Tall-Grass" Series, 142.11 of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Nebraska Sand Hills- Sand Hills Ecoregion, 44a (Chapman et al., 2001).

77. Beauty lower down- Just a short cattle walk off of the low sand dune or hummock dominated by little bluestem and upland switchgrass shown in the preceding three-slide set there was a seep formed by drainage of water from higher ground by infiltration and subsurface lateral flow (and, perhaps, some runoff of higher dunes). This seep or microenvironment of locally ponded water provided ideal habitat for bluejoint reedgrass with Brevior's or short-beaked caric sedge (Carex brevior). The wetland formed by this microrelief constituted a small "pocket" of wet prairie or, depending on interpretation, marsh.

Is there anything in God's creation that is more soothing or satisfying to the human eye than pastoral scenes such as this?

Loup County, Nebraska. Late June; early estival aspect. FRES No. 39 (Prairie Ecosystem). K-67 (Nebraska Sand Hills Prairie). Wet prairie vegetation was a variant of SRM 602 (Bluestem- Prairie Sandreed). Bluestem "Tall-Grass" Series, 142.11 of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Nebraska Sand Hills- Sand Hills Ecoregion, 44a (Chapman et al., 2001).

78. A bluejointed stand- Local population or colony of bluejoint reedgrass or, more commonly, bluejoint (Calamagrostis canadensis) in a basin or depression on tallgrass otherwise dominated by prairie sandreed, sand and/or big bluestem, switchgrass, and little bluestem. Bluejoint typically grows in wet to very mesic habitats, at least those environments that have abundant soil moisture during the growing season of this cool-season grass (a member of Aveneae, the oat tribe).

Bluejoint has a vast species range that extends from Greenland throughout the Canadian Provinces west to California and south to New Mexico and Georgia. Biological range of this "highly variable" includes northern Eurasia (Fernald, 1950, p. 157). Fernald (1950, p. 157) described four taxonomic varieties of C. canadensis. In many areas of North American rangeland bluejoint is a major forage species and one that is arch-typical of more northern prairies where it is often a major component of prairie hay.Bluejoint is widely distributed throughout more mesic range sites and microsites throughout the Nebraska SandhillsGood references for bluejoint include the standard Range Plant Handbook (Forest Service, 1940, p. G41) It was explained in the Forest Service Range Plant Handbook that bluejoint herbage is most palatable when young, which is characteristic of almost all grasses (forage in general), but grazing bluejoint at the more nutritious and palatable stages is usually problematic given the moist to wet habitats in which bluejoint grows. Livestock need to be withheld from such wet range sites until soils dry out to prevent damage to the range (ie. range readiness, the Cardinal Principle of proper season of use) and to minimize livestock, especially sheep, foot problems.

Bluejoint is a highly rhizomatous species with much of its reproduction being modular or clonal, and of a particularily opportunistic nature or growth pattern (Mcdonald and Liefers, 1993). Bluejoint was also found to be at least "semiapomictic" (Nygren, 1954; Greene, 1984). Clearly much of plant growth, including regrowth following defoliation, as well as plant regeneration in absence of shoot removal is asexual reproduction in bluejoint.

"Sourdough" is one cultivar of bluejoint that was developed and released by the Alaska Agricultural Experiment Station.with seedstocks maintained by the NRCS Alaska Plant Materials Center.

Rock County Nebraska. Late June; early inflorescence--just-emerged-from-the-boot--phenological stage.

79. Plants of bluejoint reedgrass- Several plants of bluejoint reedgrass or, simply, bluejoint on a local wet (at least seasonally) prairie in Nebraska Sandhills. Most of these polants were in earliest stages of panicle expansion (ie. just emerged from the boot; see next two slide-caption sets). Bluejoint produces robust creeping rhizomes and appears to reproduce primarily from these. Bluejoint is at least "semiapomictic" (Nygren, 1954; Greene, 1984) so that it also reproduces asexually from clonal seed. Presumedly sexual reproduction from fertilized grain also takes plaace. Bluejoint hedges it bets with regard to reproduction.

Bluejoint produces a dense sod, but the sward of this species characteristically produces "tufts" of shoots (tufted clonal units) so that bluejoint turfs give the appearance of having cespitose plants arising from an open carpet of semi-horizontal shoots (ie. bunchgrasses within a sparse sod). Such a visual impression was obvious from these two photographs.

Although bluejoint is widely distributed and an important forage grass (and with such productivity that it can become a weed) there have been remarkably few treatments of the biology of bluejoint. Sparrow and Panciera (2005) evaluated influences of mowing and nitrogen fertilizzation on bluejoint in Alaska. Their study was primarily from stanpoint of controlling bluejoint when viewed as a weed or, alternatively, use of bluejoint as a native hay crop. Another study of bluejoint in Alaska (Collins et al., 2001) involved heavy grazing in an attempt to reduce cover of bluejoint on forest clearcuts. Both of these Alaska trials found bluejoint reedgrass to be remarkably tolerant of heavy defoliation. Collins et al. (2001) attributed much of this adaptation to close cutting or grazing to the rhizomatous feature of bluejoint.

A good general reference for bluejoint, complete with reference list, was Wynia (2006).

Rock County, Nebraska (first slide) and Loup County, Nebraska (second slide). Late June; early inflorescence--just-emerged-from-the-boot--stage.

80. Business (sexually reproductive) end- Sexual shoots of bluejoint reedgrass with young panicles just about fully emerged from the boot. The individual branches of the panicles were still appressed (to a large extent anyway) against the central axes of these panicles. At this just-emerged-from-the-boot stage these flower clusters appeared to be condensed panicles, but this is not the mature, fully exerted form of this compound inflorescence which is a spreading, open panicle.

Rock County, Nebraska (first slide) and Loup County, Nebraska (second slide). Late June; early inflorescence--just-emerged-from-the-boot--stage.

81. Still unfurled- Two young panicles of bluejoint reedgrass that had just been fully exerted from their boots. The boot is the uppermost grass leaf that envelops the unemerged inflorescence and from which the inflorescence is eventually exerted when the flower cluster-bearing culm grows in length enough to be expressed through the boot. The panicle in the first photograph was not fully extended or expelled from the boot whereas the panicle in the second photograph was fully exerted from its boot at which state the boot (panicle-enveloping leaf) becomes the flag leaf. The flag leaf is the leaf subtending (immediately below) the inflorescence which, again, in bluejoint is a panicle.

These emerging and newly emerged (first and second photograph, respectively) panicles still had their branches appressed (pressed against) the central axis of the panicle. At maturity of panicle (by stage of anthesis) these branches (secondary units of the panicle) will have extended hore or less horizontally or, at least, ascendingly.

Rock County, Nebraska. Late June; early inflorescence--just-emerged-from-the-boot--stage.

82. Details at an early--though advancing--stage- Portion of a panicle of bluejoint reedgrass with branches beginnning to drop into their mature horizontal orientation (first slide) and young (immature) branches of bluejoint with branches falling down from their appressed position against the central axis revealing individual spikelets (second slide). Both panicles were still in pre-anthesis stages.

Students should take note that characteristic features of inflorescence and spikelet may be difficult to see or recognize in immature stages such as seen here.

Loup County, Nebraska. Late June; early inflorescence stage.

83. Brevor's caric sedge opr short-beaked caric sedge (Carex brevior)- Sexual shoot (first slide) and spikes in inflorescence (second slide) of Brevior's sedge over their leafy turf in a localized marsh or wet prairie in Nebraska Sandhills (two immediately preceding slides). C. brevior is in the Vignea subgenus.

Good reference for this species was Hurd et al. (1998, ps. 68-69). There is a Carex brevior group which includes (or, maybe, formerly included) C. bicknellii and/or the C. opaca complex (Rothrock and Reznicek, 2001). In other words, to rangemen of the non-taxonomists strain (which includes most of us) a bunch of these Carex species are closely related. This was shown in for Nebraska in Kaul et al. (2006, ps. 382-383) and, better yet, in the Missouri flora of Steyermark (1963, ps.340-346) under the Ovales section of Carex.

Loup County, Nebraska. Late June; fruit-ripening stage.

84. Postclimax tallgrass prairie at its Sandhills ultimate-Big bluestem-switchgrass-green needlegrass variant of tallgrass prairie. This climax range vegetation was on a moist upland above the Niobrara River. At this latitude (43 degrees north) a cool-season, festucoid mid- to tallgrass species joined the typical, panicoid tallgrasses as a local dominant to general associate species on a range in Excellent range condition class. Other grass species included sand bluestem, blue grama, needle-and-thread, and both of the major, naturalized, annual, Eurasian bromes, cheatgrass (Bromus tectorum) and Japanese chess (B. japonicus). Prairie sandreed was absent. Wild or American licorice (Glycyrrhiza lepidota), the major forb, was locally abundant.

This mixture was consistent with the observation of Clements (1920, p. 131): "Andropogon is typical of the commuity to an almost exclusive degree, but the species often mix with Stipa, Agropyron, and Koeleria to such an extent as to make the excact relationship of a particular area difficult to determine". Big bluestem, switchgrass, and Canada wildrye were included as more mesophytic of the tallgrass species as compared to little bluestem (Clements, 1920, ps. 133-134). Green needlegrass was described by Clements (1920, ps. 119, 137-138) as the more mesophytic dominant of the mixed prairie and being affilitated with Andropogon as a subclimax dominant.

Status of this range plant community as to potential natural vegetation could be regarded as ambiguous in the confusing scheme of the Clementsian monoclimax model. From the literature it appeared that this ambiguity (or downright confusion) came about as the Clementsian hierarchial scheme was modified over course of Clements' and his associates' careers. Early on, Clements (1920, ps. 131-132) interpreted this as "the subclimax prairie", the Andropogon associes", where associes was the seral equivalent of the association. "A remarkable development of this sort [the Andropogon associes] occurs in the great sandhill region of Nebraska, where Andropogon is again the dominant genus. Here the important dominants are bunch-grass, as demanded by the more rigorous water conditions, and the climax is the Stipa-Bouteloua prairie" (Clements, 1929, p. 132). Later at zenith of the Clementsian model Weaver and Clements (1938, p. 520) interpreted the "tall-grass prairie" as existing in "... three forms characterized by the same general dominants but of different ecological import". Tallgrass prairie was described as "preclimax to the forest [to the east] and may also serve as a subclimax in its development; at the same time it is a postclimax to the true prairie" The next sentence clairified the postclimax relationship to true prairie: "In this latter relation it [tallgrass prairie] is found generally in sandy plains, sand hills, and other dune areas in the grassland, the most notable being the sand-hill complex of central Nebraska" (Weaver and Clements, 1938, ps. 520-521).

In the monoclimax model a given plant community, especially a formation, could play a "double role" (has "twin identies" might have been a less confusing choice of words) being "... a preclimax to its more mesophytic neighbor and a postclimax to it more xerophytic one". The Clementsian perspective regarded units of vegetation as "but a segment of the dynamic flow of processes". Left as residual vegetation from past climates were relicts. "The great climatic changes of the past have everywhere left relict commuhities that bear either a preclimax or postclimax relation to the surrounding climaxes of the present day." "A relict is a community or fargment of one that has survived some importnt change. ... Tall bluestems (Andropogon) are relicts in the sand hills of Nebraska..." (Weaver and Clements, 1938, p. 83). "In the vast sand-hill area of Nebraska, the tall-grass postclimax attains its best development, which is assumed to relfect the climate when the prairies were occupied by the blusetems and their associates some millions of years of years ago" (Weaver and Clements, 1938, p. 86).

Nomenclatural note: at time of early to mid periods of Clements' career Indiangrass was included in Andropogon (= Andropogon nutans) so that this species was one of the '"tall bluestems" in works like Plant Indictors (Clements, 1920, ps. 132, 133).

Sandhills tallgrass prairie is postclimax as a consequence of deep sandy soil that has substantially greater water-holding capacity than what is typical of this semiarid precipitation zone. Net result is a true tallgrass prairie (a postclimax grassland) "smack dab" in middle of true or, perhaps, mixed prairie climax (the climatic or regional climax).

The big bluestem-switchgrass-green needlegrass combination presented here was the most mesic expression or variant of tallgrass prairie this author encountered in the Nebraska Sandhills excepting irrigated and subirrigated range sites or, generally, wetlands like marshes and wet prairies of prairie cordgrass (Spartina pectinata).

Ft. Niobrara National Wildlife Refuge, Cherry County, Nebraska. Late June, but cold though moist late winter-spring weather retarded phenological development of all native plant species: vernal aspect. FRES No. 39 (Prairie Ecosystem). K-67 (Nebraska Sand Hills Prairie). SRM 602 (Bluestem- Prairie Sandreed). Bluestem "Tall-Grass" Series, 142.11 of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Sandy range site. Nebraska Sand Hills- Sand Hills Ecoregion, 44a (Chapman et al., 2001).

85. Disclimax tallgrass prairie- A range plant community of red threeawn (Aristida longiseta), tumble mustard (Sisymbrium altissimum), western salsify or goatsbeard (Tragopogon dubis), wooly plantain or wooly Indianwheat (Plantago patagonia) six-weeks fescue (Festuca octoflora), cheatgrass, Japanese chess, Canada bluegrass (Poa compressa), and an unidentified Euphorbia species that developed on a black-tailed prairie dog (Cynomys ludovicianus) town. The postclimax tallgrass prairie vegetation introduced immediately above had been eliminated by overgrazing and was replaced by a lower stage that placed range shown in these two slides in Poor range condition class. The Sandhills range plant community presented in these two photographs and the single succeeding photograph was part of the same range (grazing unit or pasture) shown in the two preceding photographs that was in Excellent range condition class. There was a distance of about 150 yards between the two range plant communities with a graveled road separating the areas of Excellent and Poor range condition. Both areas of this pasture--Excellent and Poor condition-- were the same range site, Sandy. On-going overgrazing by black-tailed prairie dogs apparently was responsible for this stark contrast (again, same pasture and same range site with the two range plant communities almost conterminous with each other). This entire range (pasture) had been grazed by North American buffalo and Texas longhorns (exhibition animals for wildlife refuge visitors) in the recent past. This range was presently subject to grazing by white-teiled deer, but the only obvious or apparent cause of range degradation was that the Poor condition portion was currently a prairie dog town. For whatever reason, prairie dogs had not crossed the refuge road. Personnel working on this national wildlife refuge stated that there had not been any human manipulation of prairie dogs on this range. This was an unequivocal, unambiguous response by refuge officials in response to direct questioning by the author for whom this natural controlled experiment showed plainly the results of overgrazing by this single species of native rodent.

Black-tailed prairie dogs do not range far from their burrows. It is a well-documented.phenomenon that black-tailed prairie dogs graze closely (to a low height) range vegetation within areas of their towns. Grazing is most extreme in and around entrances to burrows where prairie dogs clip all tall plants (and most other vegetation) tlypically resulting in bare soil. This behavior results in defoliation ranging from heavy grazing to local denudation with the ultimate result frequently (even typically) being overgrazing. It was this herbivorous activity, a combination of feeding on herbage and removing plants (by clipping or biting off) that would otherwise obscure potential predators from the view of prairie dogs, that seemed--and quite obviously so--responsible for range degradation (= range deterioration) seen at wide view in these two photographs and at closer range in the next photograph. It was apparent that overgrazing by prairie dogs had converted climax (postclimax) tallgrass prairie into seral vegetation. This seral plant community was just a stage or two beyond that of the pioneer community for the sere of this range site. It was possible that larger ruminants had contributed to overgrazing and retrogression on this range, but at minimum prairie dogs had maintained the state of range depletion. The rest of this range outside of the prairie dog town had also been grazed by cattle and buffalo and that portion was in Excellent range condition class. It could only be concluded logically that black-tailed prairie dogs had induced the severe state of range retrogression shown here.

Impacts of prairie dogs, especially the more populous black-tailed prairie dog, on vegetation and other components of range ecosystems remains one of the seemingly most varied of phenomena on North American grasslands. Seemingly (or apparently) being perhaps the key to variable findings reported in the literature, scientific and otherwise. Effects--direct and/or indirect--of prairie dogs on native grazing lands have been (again, seemingly) quite difficult to "ferret out". There have been a few other topics in natural resource management within the USA that have been as open to interpretation and, therefore, as controversial as that regarding the black-tailed prairie dog and its role in ecology (and human management) of Great Plains grasslands. Affect of wolves on prey, along with reintroduction of this predator, role of fire (suppression vs. prescription), damage done by feral equids, and logging old-growth timber have garnered more press coverage but probably no more firey charged emotions among actural resource managers that this fossorial (burrowing) rodent.

Black-tailed prairie dogs have been the subject of considerable research, but the actual influence of this species on range vegetation has not been a major thrust of most these investigations. Aspects of black-tailed prairie dogs on rangelands that have been investigated was shown by the following short sampling of the range research. Vegetational changes on shortgrass range due to black-tailed prairie dogs were reported by Bonham and Lerwick (1976). Agnew et al. (1986) also evaluated shifts in plant species on mixed prairie range under grazing by black-tailed prairie dogs. Summers and Linder (1978) analyzed food habits of the black-tailed prairie dog, but did not include impacts of feeding on plant species or the range plant community. Sharps and Uresk (1990) published a literature review of black-tailed prairie dogs and associated bird and mammal species, but did not so much as mentions a single plant species. Uresk (1985) had previously analyzed changes in production of plant species on mixed prairie range in response to removal (by poisoning) of black-tailed prairie dogs. Competition between black-tailed prairie dogs and cattle was investigated by O'Meilia et al. (1982) who reported that prairie dogs reduced herbage availability but not enough to statisticlly decrease weight gains of steers. Collins et al. (1984) conducted an economic evaluation of chemical control of black-tailed prairie dog populations. They found that population reduction "was not economically feasible". Analysis of range vegetation was not an objective of this economic study.

Comparatively guaged there have been remarkabley few ecological studies, especially range experiments, that were specifically designed to analyze the role of prairie dogs on their habitat. It is remarkable that there have been so few actual investigations into this interaction. In fact, it appeared at time of this author's appraisal that the monographic investigation of Koford (1958) remained one of the few wildlife-range examinations regarding environmental impact of the black-tailed prairie dog. The more recent monograph of this species by Hoogland (1995) focused on its social life. Hoogland (1995, p. 14) devoted about two-thirds of one page (out of over 400 pages of text) to diet of black-tailed prairie dogs. This segment was a short literature review as diet was not studied in this work (other than to report on the discovery of cannibalism). Hoogan (1995, p. 22) offered this one summary sentence concerning prairie dogs and range vegetation: "By their foraging and clipping of tall vegetation, prairie dogs radically alter the plant communities of western North America". The few other investigations generally concluded that prairie dogs strongly prefer certain grass species and can reduce these (and shrubs), but on some range types such as tallgrass prairie prairie dogs may, in fact, depend on grazing by bovids (buffalo or cattle) to maintain vegetation conducive to prairie dog colonization. This suggested that prairie dogs were more a symptom or result of heavy grazing rather than a cause--at least an initial cause--of it. Reduction of heavy grazing by bovids has even resulted in elimination of some prairie dog towns as tallgrass species reclaimed seres with secondary plant succession (Koford, 1958). In contrast to the situation reported for tallgrass prairie, black-tailed prairie dogs have been observed to maintain degraded range (lower seral stages) on mixed prairie and shortgrass plains grasslands.

Foraging activity (probably feeding plus removal of plant cover for protection visibility) and burrowing by black-tailed prairie dogs decreased western wheatgrass and blue grama while increasing cover of buffalograss and forbs (Agnew etal., 1986). Uresk (1985) also reported decreased production of western wheatgrass due to black-tailed prairie dogs, but no differences in production of buffalograss or needleleaf sedge (Careex eleocharis). Bonham and Lerwick reported that black-tailed prairie dogs reduced cover of blue grama, increased cover of buffalograss and sixweek's fescue, had no effect on needleleaf sedge, and increased number of plant species (both annual and perennial).

Some of the more recent studies reported in the peer-reviewed literature seemed to be the equivalent of "political correctness" for rodents. It seemed to this author that many findings were couched in equivocated, obtuse wording so as to suggest unnecessarily vague conclusions, if conclusions were reached at all. For example, some statements were made in regard to "grass" or "forbs" without being specific as to species or successional status (eg. no distinction as to decreaser, increaser, invader; not specific as to which seral stages certain species belonged). In one notarious instance, an informant very familar with a specific study explained--off the record and only with assurances of ambiguity--to this author how one trial was biased from the onset by prejudiced investigators and the data "cooked" to make it seem that black-tailed dogs had beneficial impacts on mixed prairie range that they had degraded by overgrazing. This "research" was eventually reported in a major journal devoted to grazing lands.

Prairie dogs influence on range has been discussed in most of the standard textbooks (Stoddart and Smith, 1955, ps. 201-202; Humphrey,1962, ps. 127-129; Holechek et al., 2004, ps. 486-487). All seem to have concluded the obvious. These native rodents (there are four prairie dog species in North America) have been an integral and major component of natural range ecosystems for millenia so that they could not be responsible for destruction of these grazing lands (or at least not damage from which ranges could not recover). At the same time, control of prairie dog populations (reduction of prairie dog numbers) has been a successful range improvement practice. Holechek et al. (2004, p. 487) summarized that prairie dog control programs "... have undoubted imporved range condition and increased the amount of forage available to livestock", yet there is a net benefit to maintaining some prairie dog towns. No knowledgable, sane person would advocate extermination of prairie dog species. Obviously, all native species are protected by the Endangered Species Act, a federal law strongly upheld by the United States Supreme Court.

86. Disclimax up close- A postclimax tallgrass prairie range degraded by black-tailed prairie dogs to a seral stage community of red threeawn, naturalized Eurasian annual forbs and annual grasses. Major plant species visible in this "photoquadrant" besides red threeawn were tumble mustard and wooly plantain or wooly Indianwheat . Land area in this photograph was part of a black-tailed prairie dog town. This slide provided a closer-in view of the plant community of the depleted range presented in the two preceding slides. Featured in this photograph was the den or nest site (mound and cleared area around it) of western harvester ant (Pogonomyrmex occidentalis). Like weedy plant species this hymenopteran was also more common on the depleted (seral) part than on the climax portions of this range which was on a moist sandy upland above the Niobrara River. Both the degraded grassland ( the prairie dog town) and pristine portions of grassland were on the same range and same range site (Sandy). The big bluestem-switchgrass-green needlegrass climax tallgrass prairie, the potential natural range vegetation, was shown and described in the two-slide and caption set that preceded the two photographs and lengthy caption immediately above.

This range had been grazed by North American buffalo and Texas longhorns a few years prior to time of these photographs. These larger ruminants had been removed and the range was grazed only by black-tailed prairie dogs and other small mammals, insects, birds, and, sometimes (at least available to) white-tailed deer. It is possible that bison and cattle had contributed to the obvious overgrazing and depletion of this range, but the state of degradation had been maintained by prairie dog action. Plus, as was explained in the preceding caption, this entire range had been grazed by buffalo and cattle with the portion not in prairie dog town being climax big bluestem-switchgrass-green needlegrass tallgrass prairie (ie. in Excellent range condition class).

Dominance by red threeawn was likely explained by the finding that black-tailed prairie dogs avoid eating Aristida species even when these are important grasses on prairie dog towns (Summers and Linder, 1978). Perennial threeawns are invaders on this tallgrass plant community.

Proper degree of use of climax range plants had been exceeded on the prairie dog town. This is the state of defoliation recognized as overuse. Extended overuse eventually leads to overgrazing. Prolonged overuse (= overgrazing) results in changes to the range plant community with species of lower succsssional state replacing plant species of higher succcessional order. Replacement of big bluestem, switchgrass, and green needlegrass by red threeawn and naturalized Eurasian weeds were changes on the prairie dog town featured here. This dynamic in range plant species is part of the process along the range sere that is called retrogression. Retrogression constitutes the process of range deterioration; stated more specifically perhaps, range deterioration is the outcome of the process of retrogression.

Successional sttatus of the range plant community is determined (largely as species composition) and designated as range condition class. Range condition (class) is the successional state of the range vegetation compared to climax or potential natural vegetation at a point in time. Changes in in range condition over time is range trend. Downward range trend is the measured result (outcome) 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 (mismanagemenrt) 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.

This tallgrass prairie range depleted to a seral state of lowest-stage perennial, native annuals, and Eurasian weeds (annual grasses and forbs)--all species of which are ecological invaders--was about as extreme a case of overgrazing-induced retrogression as could exist for thiis range site except for the pioneer (colonizing) stage.

87. All-in-one composite view- Landscape-scale view of the Nebraska Sandhills showing the grandure of this landform with its various range types and range sites. Range in the foreground and distant background was primarily that of a Sands range site with an amazing species diversity with such major grasses as little and sand bluestems, prairie sandreed, needle-and-thread, sand lovegrass, sandhills muhly, sand dropseed, blue and sideoats gramas, and Junegrass. On this sixth full day of spring (time of photograph) and after a wet, cold late winter and spring needle-and-thread was more conspicuous--a seasonal aspect dominance--than would be the case later when the warm-season grasses (almost all other grass species on this range) reach maturity (peak herbage yield). Typically, Junegrass was present but not particularly common (frequent).

The wetland in mid- to near background was a freshwater marsh of tule or bulrush species, spikerush, and common reed. Nebraska Sandhlls marsh vegetation was treated immediately below.

Cherry County, Nebraska. Late June; early estival aspect. Tallgrass prairie was FRES No. 39 (Prairie Ecosystem). K-67 (Nebraska Sand Hills Prairie). SRM 602 (Bluestem- Prairie Sandreed). Bluestem "Tall-Grass" Series, 142.11 of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Freshwater marsh was FRES No. 41 (Wet Grasslands Ecosystem). K-42 (Tule Marshes). No SRM in Northern Great Plains chapter (Shiflet, 1994) for marsh. Upland tallgrass prairie was Sands range site; marsh was Wet Subirrigated to Wetland range sites (Natural Resources Conservation Service, 2005). Nebraska Sand Hills- Sand Hills Ecoregion, 44a (Chapman et al., 2001).

88. Growing on a subsurface layer of water- Subirrigated meadow--probably should be interpreted as a marsh--in Nebraska Sandhills. Bluejoint reedgrass appeared to be the climax dominant, but introduced agronomic and weed species had displaced much of the original bluejoint. Instead the human-modified range plant community had become a "man-made mixture" of two Eurasian grasses, redtop (Agrostis alba= A. stolonifer) and creeping foxtail (Alopecurus arundinaceus); the native common spikerush (Eleocharis palustris), and the Eurasian weedy grasslike plant, round-fruit rush (Juncus compressus).

Note on organization: redtop was treated in Introduced Forages, Grasslands portion, herein.

Subirrigated referred to the condition in which the surface aquifer was high enough in the soil profile or regolith that roots of plants growing on the land extended down into this wet zone. Plants are not necessarily those species that are obligate or facultative phreatophytes, but instead were plants with typical rooting depth which was adequate to absorb water from the shallow surface aquifer. As was described above, ponded water or shallow, subsurface soil water in the Nebraska Sandhills are results from various conditions of soil texture, microrelief, and related hydric phenomena (subsurface lateral flow, deep percolation, overland flow, etc.).

Cherry County, Nebraska. Mid-June; late vernal aspect. FRES No. 39 (Prairie Ecosystem). K-67 (Nebraska Sand Hills Prairie). Wet prairie vegetation was a variant of SRM 602 (Bluestem- Prairie Sandreed). Bluestem "Tall-Grass" Series, 142.11 of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Subirrigated range site (NRCS, 2005). Nebraska Sand Hills- Sand Hills Ecoregion, 44a (Chapman et al., 2001).

89. Subirrigated sward- Turf of a wet prairie or marsh (as the case may be) of bluejoint reedgrass, redtop, creeping foxtail, common spikerush, round-fruit rush, and similar graminoid species. There were even a few plants of bird'sfoot trefoil (Lotus corniculatus), an agronomic Eurasian papilionaceous legume.

Note or arrangement: bird'sfoot trefoil was included in the chapter, Introduced Forages in the Grassland section of Range Types.

90. Common where it was subirrigated- Common spikerush with some round-fruit rush comprised the range vegetation presented in this "photoquadrant" from a subirrigated wet meadow (marsh or wet prairie, depending on interpretation and/or definition) in the Nebraska Sandhills. View of this entire, subirrigated, wetland range plant community was presented two slide-caption sets above.

Cherry County, Nebraska. Mid-June.

91. Spiked where subirrigated-Common or creeping spikerush (Eleocharis palustris= E. macrostachya) on a subirrigated wet meadow (or marsh). First slide presented a number of sexual shoots while the second slide showed details of two inflorescences (spikes) at closer distance. Spikes were in anthesis. This species (or whatever the taxon as the case may be) has been a "spike in the side" of taxonomists. E. palustris is a taxonomic complex (Kaul et al., 2006, ps. 413-414) including E. macrtostachya and E. smallii recognized by Great Plains Flora Association (1986).

Good--though brief--treatment of this wetland grassland plant was by Correll and Correll (1975, ps. 381-382). The author could not find any discussion of Eleocharis species in regards to their forage (feed) value, but his observations were that these plants are eaten only infrequently under cattle grazing. Their main economic value would appear to be protection and formation (building) of soil on erosion-prone wetlands. Eleocharis species are commonly major--sometimes sole--plant species on marshes and related wetlands.

Cherry County, Nebraska. Mid-June; anthesis.

92. As if sand wasn't scourer enough- Common scouring rush (Equisetum hyemale) on subirrigated meadow, a freshwater marsh converted into a semidomesticated wetland pasture of naturalized introduced forage species. This pterophyte--a vascular, spore-producing plant--was a neighbor of common spikerush, a native graminoid, and round-fruit rush, a Eurasian weedy grasslike plant. All of these species were much less abundant and important than redtop and creeping foxtail which are also naturalized introduced species.

The sexual (spore-bearing) shoots in these two slides had strobili (cones) at their apex. These are, of course, the sporophytes, the sporophytic generation. The poka-dot appearing spots on these cones are sporophylls which terminate with disk bearing sporangiophores.

The name "scouring rush" comes from the pioneer practice of using shoots of this (and other Equisetum) species to clean (scour out) cast iron cookware and pewter eatingware. This Equisetum species does not grow lateral, leafy shoots (leaf whorls) in the same manner as common horsetail, (E. arvense) so this morphological feature combined with the practice of using its shoots for scouring utensiles makes "scouring rush" a more fitting name (even though Equisetum species are not rushes). The scouring capacity of the horsetails derives from the impregnated silica that is deposited and arranged in ridges along the shoot.

The timeless Notes on Western Forbs (Dayton, 1960, ps. 3-5) was recommended to readers. This author indicated that E. hyemale was used for scouring floors and that its evergreen habit (even in cold winters) was basis of the specific epithet hyemale referring to winter. Good references for basic biology of Equisetum species were Eames (1936, ps. 88-115) and Holman and Robbins (1946, ps. 516-522).

Cherry County, Nebraska. Mid-June.

93. Lake's edge and "rings" of marsh vegetation - Wetland range vegetation at perimeter of a freshwater lake in Nebraska Sandhills. There were two major range plant communities in this marsh range both of which were local zonal communities arranged spatially as more-or-less concentric "rings" around and along the lake. The outermost concentric-patterned community was dominated by narrowleaf (= narrow-leafed or narrow-leaved) cattail (Typha angustifolia), sometimes exclusively as a consocies (seral equivalent of a consociation) and at other times growing with giant or late goldenrod (Solidago gigantea) with this latter species varying from associate to co-dominant. Inward from this was a "ring" (the innermost concentric zone) dominated (usually solely populated) by three-square (sometimes unhyphenated as threesquare) or chairmakers' rush or bulrush (Scirpus americanus= S. pungens= Schoenoplectus pungens) which was also a consocies (Clements, 1936, p. 264).

These two photographs presented the focal perspective (angle of view) across the two plant communities (ie. looking into and/or directly over top of the zones of wetland vegetation). Said another way, it was a tall view across (= at roughly a right angle to) the concentric zones ("rings") of narrowleaf cattail and chairmakers' or three-square bulrush (roremost, most of foreground, and hindmost, most of background, respectively; especially so in first photograph). The perspective or view "in line" (straight) with the zones ("rings") was presented in the photograph immediately theser two slides.

The"bed" (stand) of cattail and bulrush or tule could be regarded as a population (ie. a colony) or, alternatively, as a consocies. Traditionally (historically) herbaceous wetland vegetation (marshes) comprised of tule or bulrush, cattail, and/or common rees (Phragmites communis) marshes were included as part of the "reed swamp" and interpreted as seral stages along hydroseres or alternatively as subclimaxes or disturbance (by flooding) climaxes (disclimaxes) in monoclimax theory (Weaver and Clements, 1938, ps. 63, 100, 454) or as an edaphic climax in polyclimax theory (Tansley, 1923, ps. 31, 33, 34, 59) though Tansley sometimes interpreted these as Clementsians consociations which implied climax (climatic climax) status. Clements (1936, p. 264) specified that the "reed-swamp" was "the univrsal example" of a special form of subclimax, the serclimax, "a seral community usually one or two stages before the subclimax". This was how "red-swamp" was described in Weaver and Clements (1938, p. 63), but without using this term of famed (or infamous) Clementsian terminology (doctrine).

Cherry County, Nebraska. Late June; early estival aspect. FRES No. 41 (Wet Grasslands Ecosystem). K-42 (Tule Marshes). No SRM in Northern Great Plains chapter (Shiflet, 1994) for marsh. In Brown et al. (1998, p. 45) would be Bulrush Series 242.33 ( none was shown, but the vegetation obviously does exist) of Plains Interior Marshland 242.3. Wet Subirrigated to Wetland range sites (NRCS, 2005). Nebraska Sand Hills- Lakes Area Ecoregion, 44d (Chapman et al., 2001).

94. "Rings" around a marsh- Zonation of wetland vegetation around margins of a freshwater lake in the Nebraska Sandhills. Concentric zones at local scale of narrowleaf (= narrow-leafed or narrow-leaved) cattail (the outer or exterior "ring" of plant life) and three-square bulrush or chairmakers' rush (the inner or interior "ring" of plant life) that developed at different linear distances from the outer zone of a freshwater lake. This was another view of the same lake and the two prominent plant commuities of marsh vegetation introduced in the immediately preceding two photographs. There was some naturalized smooth brome as well as some plants of giant or late goldenrod in the cattail-dominated community.

This photograph presented this wetland range vegetation from an "in-line" perspective, the focal view looking from the end of the zones rather than from the middle of (looking across) the "rings" of the two prominent plant communities. In other words, this photograph gave a more-or-less "end on" look or view in contrast to the "through the middle" perspective provided by the two immediately preceding photographs. Or, In farm boy parlance, this slide gave a "gander down or between the rows" whereas the two preceding photographs were "look-sees through or into the rows".

Both the cattail and three or chair-makers' bulrush zones or concentric stands where the potential natural vegetation, but in the Clementsian monoclimax model they were long-persisting seral stages on a hydrosere and thus consocies rather than consociations of a climax. In the Tansian polyclimax model these two stands were edaphic (or, perhaps more precisely, hydric) climaxes. Most specifically from the interrpretation of monoclimax theory these two local, zonal, wetland plant communities were each a serclimax, "a seral community usually one or two stages before the subclimax" (Clements, 1936, p. 264).

95. Stand of narrowleaf cattail- Outer edge of a zonal community, a consocies or edaphic (or hydric) climax in monoclimax or polyclimax theory, respectively, dominated by Typha angustifolia growing around a freshwater lake in theNebraska Sandhills. Late or giant goldenrod was a local associate or, sometimes, co-dominant with narrowleaf cattail. The inner zonal plant community (interior "ring" of vegetation or plant life) of this sandhills marsh was a stand (consocies or edaphic-hydric climax) of chair makers' (sometimes shown hyphenated or as one word) or three-square bulrush (=tule). Part of the stand of chair-maker's tule was visible in midground.

This photograph furnished a "straight through" and/or "right across the top" view of this sandhills marsh vegetation.

96. Stand of threesquare (sometimes hyphenated) or chairmakers' rush or bulrush (Scirpus americanus= S. pungens= Schoenoplectus pungens)- Stand of three-square bulrush at edge of freshwater marsh in Nebraska Sandhills. In many parts of the west another name for bulrush is tule. Three-square tule is one of the more widely distributed Scirpus species, none of which typically receive much grazing. In this stand there were many dead shoots from previous years. Some of the live shoots around this Sandhills lake had inflorescences (see next slide), but most did not.

Note on "musical chairs" nomenclature: For generations the various species of bulrush or tules where designted as being in the genus, Scirpus. Recent taxaonomic treatments have split the traditional Scirpus into several genera (including Schoenoplectus). The author of Range Types has pointed out repeatedly that the older and more thorough descriptions of native vegetation used the then current binominals so that readers must confer both the older and the recent taxonomic literature in order to be scientifically "bilingual" ("tri-", "quadra-", etc.) in order to interpret the seminal (and classic) descriptive works.

Cherry County, Nebraska. Late June (near peak biomass, some shoots at full-boom).

97. Flower cluster of threesquare or chairmakers' bulrush- An inflorescence cut from a shoot of Scirpus pungens (= S. americanus) in the stand at edge of the freshwater lake (a freshwater marsh) presented above. Wind speeds up to 60 miles per hour on this day necessitated photographing this flower cluster in a sheltered area off of the bulrush shoot.

Cherry County, Nebraska. Late June (spikelets were just past anthesis).

98. Different version of a Nebraska Sandhills marsh- Marsh community around an ephemeral lake dominated by hardstem bulrush (Scirpus acutus) and broadleaf cattail (Typha latifolia). These two species of grasslike plants were co-dominants of the upper layer of vegetation. Main difference between this marsh vegetation and that treated immediately above was in Scirpus and Typha species. Otherwise physiogonomy, structure, and so on were the same (or "purt nar' so").

Successional status of wetlands such as Sandhills marshes historically was interpreted differently depending on which climax theory--monoclimax, polyclimax, or climax pattern--was followed. Interpretation from Clementsian monoclimax theory and Tanslian polyclimax theory was used in ediscussion of the preceding Typha angustifolia-Scirpus americanus (= S. pungens= Schoenoplectus pungens) marsh. There was also a concluding summary at end of the section on marshes (below). )

Comparison among these and other marshes in the Sandhills demonstrated dramatically the biodiversity and ecosystem variantion within relatively short distances of this remarkable assemblage of range plant communities.

99. Now for the lower layer- Small duckweed (Lemna minor) formed a water-surface layer on the freshwater marsh with a Scirpus acutus-Typha latifolia co-dominant upper herbaceous layer. This would be a nice habitat for ducks to stop over during migrations (although duckweed and other green vegetative might be gone by that time).

100. Hardstem but not hard to identify- Sexual shoots, including inflorescence of hardstem or great bulrush (Scirpus acutus= S. pungens= Schoenoplectus pungens) growing on a Nebraska Sandhills marsh (the one presented shortly above). This is one of the most readily identifiable of the Scirpus species because the large needle-like apex of the shoot (second of the two slides especially) is prominent and indisputable. (It was much more difficult to get a crisp photgraph of this subject given need for slow shutter speed due to slow fiilm in conjuction with a strong Sandhills wind. God provided momentary pauses between gusts that permitted images presented here.)

The common name, great bulsush, is as aptly descriptive as the specific epithetic, acutus, given the "great" size of this plant. Some of the shoots shown in above photographs reached over ten feet in height. Unfortunately, other Scirpus species are also known as great bulrush, including S. tabernaemontani and S. validus. S. At any rate, Scirpus acutus is one of the larger herbaceous wetland plants in the Nebraska Sandhills.

Good source for brief discussion of this species was Correll and Correll (1975, ps. 360-363 passim). Common spikerush (Eleocharis palustris= E. macrostachya) Correll and Correll (1975, ps. 381-382) E. paalustris is a taxonomic complex (Kaul et al., 2006, ps. 413-414) including E. macrtostachya and E. smallii recognized by McGregor et al. (1986).

Cherry County, Nebraska. Late June;anthesis).

101. Harder looks- Two closer-in views of shoot (stem) details and inflorescences of hardstem bulrush in a Nebraska Sandhills marsh. As was the case for the two preeding photographs these two images were of shoots produced in the freshwater marsh featured above.

There are remarkably few publications devoted to the bulrushs (or, for that matter, other wetland grasslike plants) other than brief taxonomic treatments. One recommended reference fo Scirpus acutus was the NRCS Plant Guide (Tilley et al, 2011)

Cherry County, Nebraska. Late June;anthesis).

102. Basis of the common name- Shoot bases of hardstem bulrush (Scirpus acutus) in a marsh in Nebraska Sandhills. These shoots were were some of the same ones whose apices, including inflorescences, were shown in the two immediately preceding slide-caption sets. It is commonly accepted by keen observers that livestock and big game make little use of bulrush shoots. Furbearers like beaver (Castor canadensis) and muskrat (Ondatra zibethicus) frequently consume rhizomes of Scirpus species and use their shoots in construction of lodges or, in case of beaver, somewhat in dam-building.

There are a number of ecotypes of S. acutus some of which were released as selections by Aberdeen Idaho Plant Materials Centers (Tilley et al, 2011).

Cherry County, Nebraska. Late June;anthesis).

Taxonomic reminder: It was explained above in this chapter that some authors have split the traditional Scirpus genus into other genera including Schoenoplectus which--as this was written--was the current nomenclatural rage for some of the traditional Scirpus species including S. acutus. Obviously this author rejected the new nomenclature, but did acknowledge it to minimize confusion for readers of Range Types of North America.

103. Freshwater marsh seen from higher ground- The narrowleaf cattail-chair makers' bulrush marsh introduced and described above was shown here at a slightly higher elevation and farther distance from shore of a freshwater lake in the Nebraska Sandhills. At this upper- and outermost margin of the lake wetland there was a partial, circular zone (an irregular "ring") of marsh vegetation dominated by late or giant goldenrod. This large, herbaceous composite was an associate or, rarely, a co-dominant with narrowleaf cattail in a zone of wetland vegetation adjacent to and outward (farther from the lake shore) from an inner "circle" of three-square or chair makers' rush. Unlike the "pretty much" continuous, circular-shaped stands (consocies or edaphic-hydric climaxes) of the grasslike range plants, the wetland plant community dominated by late or giant goldenrod was disocntinuous with the cattail-dominated zone which was wider at some locations so as to either 1) force out (out-compete or "overpower") goldenrod or 2) limit this large forb to co-dominant or, more commonly, associate species status.

The clumps of grass (obvious as last year's dry, weathered shoots) in the giant goldenrod-dominated zonal community was bottomland switchgrass.

The zonal range vegetation along the shore of this freshwater lake (visible in midground as a dark or gray-green "fiinge" or "border" aound the lake) was chair makers' or three-square rush (bulrush).

104. * "Reed-swamp" in Nebraska Sandhills- Freshwater to brackish marsh on a mosaic of range sites (Wetland through Wet Subirrigated and even some isolated spots of Subirrigated) present as a patchwork of several range plant communities. The outermost (least hydric) zone or range vegetation was dominated by common reed (Phragmites communis= P. australis). Common reed was accompanied by two species of large, rank-growing forbs. One of these forabs was giant or late goldenrod. The other big "weed" or "wild flower" could not be identified by this photographer in its vegetative stage. There were also trace quantites of naturalized smooth brome in this outermost stand of marsh vegetation.

The zonal or "ring" (circle-shaped spatial pattern) or wetland vegetation interior to that of common reed was narrowleaf cattail while that which was interior to the circular stand of cattail was threesquare or chair-makers' bulrush (tule). These two zonal range plant communities were presented above where they were described and interpreted as to successional status.

The interior of this marsh was somewhat to semi-saline with this brackish habitat comprised almost (essentially) of inland saltgrass. This chloridoid grass was shown in greater detail and described at various other "stops along the way" in Range Types, including above in this chapter (the section that covered wet prairie on floodplains of the Salt Fork of Arkansas River (Great Salt Plains National Wildlife Refuge).

* "Reed-swamp" is a historic designation for a marsh dominated by such wetland plants as common reed (an arundinoid grass), cattail (a grasslike plant in its own family, Typhaceae), and various members of the sedge family (Cyperaceae) including bulrushes, caric sedges, umbrella sedges, and spikerushes. In modern and now standardized (more or less) classifictions of plant communities the term swamp is restricted to wetland vegetation dominated by woody plants, especially trees, whereas wetland vegetation dominated by herbaceous species, such as grasses and grasslike plants, is now precisely designated as marsh. This speificity in use of these particular terms was not a convention and nearly universal standard at time of the first (founding) generation of Anglo-American plant ecologists. F.E. Clements and his students--for all of their otherwise precise usage of ecological terms and ultra-specific, term-rich lexicon--used "swamp" as a generic term for wetland. Hence, "reed-swamp" was applied to some of the exact kinds of wetland vegetation (Clements, 1916, ps. 188, 198, 244; Clements, 1936, p. 264; Weaver and Clements, 1938, ps. 62-63, 77, 100, 404, 454) that were presented in this section on the Nebraska Sandhills.Clements' close friend and British colleague, Sir Arthur Tansley, also relied on the designation of "reed-swamp" for the same herbaceous wetland vegetation (Tansley, 1926, ps. 31, 33, 34, 59-60) as did the more recent (less dated) British plant geographer (Polunin, 1960, ps. 314, 319, 325, 326, 372, 373, 405, 461, 463, 505).

Some of the genera and even species of the "reed-swamp" are nearly global in their distributions although these are typically discontinuous ranges, especially of the diffuse pattern or type (Polunin, 1960, 188). For example, common reed (various subspecies, ecotypes, or whatever of Phragmites communis)has traditionally been cited as "the most widely distribueedd vascular plant species in the world" (Polunin, 1960, p. 98).

105. Edge of the bed- Edge or margin of a reedbed (stand of wetland range vegetation dominated by common reed) with sensitive or bead fern (Onoclea sensibilis) as associate (or maybe both species were co-dominants), far right side of first slide and right half of second slide, that adjoined another wetland range plant community of various graminoids, an unidentifiable Solidago species, and diamond or Missouri willow (Salix eriocephala), left three-fourths of first slide and left half of second slide. In the lower herbaceous layer, especially of the reedbed with sensitive fern, there were numerous plants of a Polygonum species that could not be identified in their early stage of vegetative growth.

Note that common reed was gradually invading the other vegetational zone (lower wetland plant community to adjacent left of reedbed). This was most evident in the second slide.

Valentine National Wildlife, Cherry County, Nebraska. Late June; early estival aspect. FRES No. 41 (Wet Grasslands Ecosystem). Within Cold Temperte Marshland 242, Plains Interior Marshland 242.3, this would be a Giant Reed Series 242.33 of Brown et al. (1998. p. 45) except that Giant Reed was shown only for marsh vegetation in more southern latitudes (Brown et al., 1998, p. 47). Wet Subirrigated to Wetland range sites (NRCS, 2005). Nebraska Sand Hills- Lakes Area Ecoregion, 44d (Chapman et al., 2001).

106. Interior of reedbed in Nebraska Sandhills- Marsh in which sparsely populated shoots of common reed formed an upper vegetational layer while a denser population of shoots of sensitive or bead fern with fewer shoots of some knotweed or smartweed (Polygonum sp.) comprised a lower herbaceous layer. This wetland vegetation was a form of the widely distributed and frequently described "reedbed" ("reed-bed").

107. Sensitive to range forbs- Local population of bead or sensitive fern (Onoclea sensibilis) as a lower herbaceous layer beneth an upper layer of common reed. Stalks of bead fern with their spore-bearing structures were visible. It was obvious where the common name of bead fern was derived.

Valentine National Wildlife, Cherry County, Nebraska. Late June.

108. Another subtype, variant, or form of Nebraska Sandhills marsh- Introduced in these two slides and further shown and described in more detail in the next two-slide sets was another caattail-tule marsh that had developed in the far western extremities of the Nebraska Sandhills. This herbaceous, wetland vegetation was almost exclusively monocotyledonous in species composition, structure, and physiogonomy.

General appearance of sandhills marsh range with emphasis on physiogonomy and structure of vegetation along with distinct zonation and the mosaic of wetland plant communities. The two major range plant communities of this marsh were upright communities. The tallest, linear-leaved stand of vegetation (midground in both slides) was a combination of broadleaf (alternatively, broadleaved or broadleafed; with or without hyphens) cattail (Typha latifolia), hardstem bulrush (Scirpus acutus= Schoenoplectus acutus), and softstem bulrush (Scirpus validus= S. tabernaemontani= Schoenoplectus validus= S. tabernaemontani). Some dead stalks of this cattail-bulsrush vegetation was in foreground of first photograph.

The second upright plant community of this marsh was the lower-growing (with lighter-, brighter-green coloration) range vegetation in foreground of both photographs. This lower, upright community was co-dominated by broadleaf arrowhead (Sagittaria latifolia) and an of unidentified species of spikerush (Eleocharis sp.) Not visible in these photographs and present as widely scattered, small patches floating on surface of still water was common or little duckweed (Lemna minor). Species of algae were associated with duckweed. These species comprised a floating plant community that was undeveloped at this early summer season.

Both of the major (the upright) wetland plant communities grew as zonal vegetation occurring as linear or beltlike stands arranged according to depth of water. The arrowhead-spikerush community was limited to environments of moist to wet mud and shallow water (one to three inch depths). The cattail-bulrush community grew in deeper water (ie. as emergents). In some local areas these upright, range plant communities occurred as continuous zones whereas in other locations they were present in an interrupted linear spatial araangement.

109. Nebraska Sandhills wetland- Progressively closer views of the tall, upright marsh community (cattail-bulrush zone) in backgrounds and short, upright marsh community (arrowhead-spikerush zone) in foregrounds that were introduced in the preceding two-slide set. Little or small duckweed and algae were present to form a floating wetland community that was still relatively undevelopd and very restricted at this early summer portion of the warm-growing season. This floating plant community was not visible in these photographs.

None of these plant species were flowering at time of photographs. A drought of five to six years duration had preceded this year's growth of plants.

110. Marsh smack dab in Nebraska Sandhills- A general view (first photograph) followed by a close-in, detailed look (second photograph) of a broadleaf cattail-hardstem bulrush-softstem bulrush wetland plant community in a marsh at the western margin of the Nebraska Sandhills. This tall, upright, emergent zone of marsh vegetation grew in and at outer edge of deeper water whereas a zone of arrowhead-spikerush comprised a lower, upright vegetational zone limited to wet mud or land covered only with shallow water (depths of roughly one to three inches). This latter range plant community was not shown in either of these two photographs (instead refer back to immediately preceding slides).

Widely scattered and small patches of little duckweed and algae that were still in early stages of seasonal development made up a floating wetland plant community in this marsh. This third--and minor at this stage of the warm-growing season--marsh community was barely discernable in the second photograph.

The gently rolling dunes of a Sands range site were shown in the background of the first slide as evidence that this marsh, this range wetland, was indeed in the Nebraska Sandhills.

111. Wetland in the western Nebraska Sandhills- A freshwater marsh had formed near the westernmost boundary of the Nebraska Sandhills. This was in the area where mixed prairie gives way to the postclimax tallgrass prairie. Some of the herbaceous species, including the major grasss were naturalized agronomic species including redtop (Agrostis alba) and reed canarygrass (Phalaris arundinacea). The dominant herbaceous species of this marsh was broadleaf cattail (Typha latifolia). The first photograph presented the agronomic forage grasses while the second photograph showed an example of a cattail colony (right margin foreground to left background). Other herbaceous species included a spikerush (Eleocharis sp.) and marsh muhly (Muhlenbergia racemosa). Marsy muhly was still in rather early growth, but was prominent in central foreground to midground (largely as dead, weathered shoots) in the second slide.

Trees of peachleaf willow (Salix amygdaloides) and plains cottonwood (Populis deltoides var. occidentalis) were growing in an interrupted spatial pattern or dispersion along margins of the wetland. .

112. Cattail colony with lesser associates- Stand of broadleaf cattail in a freshwater marsh in the western Nebraska Sandhills. In contact with cattails on outer edges of the marsh were miscellaneous forbs including unidentified Solidago sp(p). plus naturalized reed canarygrass and redtop. Trees on perimeter of this wetland were plains cottonwood and peachleaf willow.

113. Woody wetland on the prairie- No, this is clearly not tallgrass prairie or, for that matter, grassland of any range type. This is an example of a variant of the cottonwood-willow forest cover type (SAF 235; Eyre, 1980, p. 113) which was covered separately under Rocky Mountain Riparian and Related Wetland Range section of the Willow & Riparian chapter of (under) Shrublands. This woody plant-dominated wetland was included here under the postclimax tallgrass prairie of the Nebraska Sandhills because it 1) is part of the range vegetation of the Nebraska Sandhills, 2) was conterminous with the herbaceous wetland vegetation (mostly freshwater marrsh) presented immediately above, 3) was consistent with eastern cottonwood and willow plant communities in other tallgrass regions such as those in the Flint Hills, tand 4) was needed to show the continuum of range vegetation found within major grassland regions.

The wetland scrub in foreground was a combination primarily of sandbar or coyote willow (Salix exigua) and some peachleaf willow (S. amygdaloides) which was contiguous (in contact) with the freshwater marsh vegetation (the Typha-Scirpus-Sagittaria wetland community) presented immediately above. Trees in background were peachleaf willow and, the larger trees, plains eastern cottonwood (western variety; .Populis deltoides var. occidentalis).

Leaves, twigs, and bark are (at least, can be) valuable browse for livestock (including cattle and hroses) and especially for wildlife species varying from cervids like deer to beaver (bark in particluar).

Cherry County, Nebraska. Late June; early estival aspect. FRES No. 17 (Elm-Ash-Cottonwood Forest and Woodland Ecosystem). K-89 (Northern Flood-plain Forest) was closest though certainly not a "perfect match". SAF 235 (Cottonwood-Willow). Plains and Great Basin Riparian Scrub biotic community, 232.2- Willow Series, 232.21 (Brown et al., 1998, p. 44). Part of Wet Subirrigated to Wetland range sites (NRCS, 2005). Nebraska Sand Hills- Sand Hills Ecoregion, 44a (Chapman et al., 2001).

Successional summary of marshes and related wetlands: In the monoclimax theory Weaver and Clements (1938, p. 62-63) interpreted Scirpus-Typha-Phragmites as the "reed-swamp stage" of a hydrosere that would (again, in theory) end in a climax forest or grassland. In polyclimax theory, this wetland plant community most logically would be interpreted as a topographic or, somewhat less logically, an edaphic climax. Details of these successional views was presented above in the first caption devoted to a Typha-Scirpus marsh.

114. Prairie sandreed (Calamovilfa longifolia)- This attractive tallgrass is the dominant species on deep sand sites in the greater Sandhills Region of Nebraska and adjoining areas of Colorado, Kansas, and South Dakota. This is locally a major hay grass and, unlike associated panicoid grasses such as the bluestems, prairie sandreed "cures on the vine" (ie. retains higher nutritive value in dead herbage) much like other eragrostoid species such as the less productive grama grasses and buffalograss. Prairie sandreed is often the key species on many range sites (and ranches).

Hays County, Nebraska. July.

115. Prairie sandreed with phyto-friends- Local colony of prairie sandreed with some companion forbs on plains cottonwood-dominated bottomland range on including prairie sagewort (Artemisia frigida), western ragweed (Ambrosia psilostachya), prairie coneflower or Mexican hat (Ratibida columnifera); annual sunflower (Helianthus annus), four o'clock (Mirabilis linearis), and wild licorice (Glycyrrhiza lepidota). The second (vertical) slide was a closer-in, more detailed view of the left foreground of the first slide. This second photograph presented a better view of shoots of prairie sandreed.

This local assemblage of range plants was on the floodplain of an interrupted or intermittent stream (whichever it was) in the semiarid zone of the Colorado Piedmont. .

East Bijou Creek, Elbert County, Colorado. Mid-July; onset of anthesis.

116. At base- Basal portion of shoot of prairie sandreed with parts of sheath and blade of the four lowermost leaves. More details were presented in the next subsequent slide. This shoot was growing on East Bijou Creek, an intermittent or interrupted (whichever one), in the Colorado Plateau, the physiographic unit just west of the High Plains or Llano Estacho (Staked Plains) of the Great Plains province.

Elbert County, Colorado. Mid-July.

117. Unit of a reed- Phytomer--fundamental, repeating unit of a grass shoot including upper half of a node, internode, and lower half of next highest node complete with leaf--of prairie sandreed (first slide) and details of culm, sheath, and colar of this phytomer (second slide). Floodplain of East Bijou Creek, an interrupted or, maybe, an intermittent stream in the semiarid tableland of the Colorado Piedmont.

Elbert County, Colorado. Mid-July.

118. Atop the reed of a reed- The panicle--the entire structure--of prairie sandreed (first slide), upper or distal portion of this panicle (second slide), and one branch of this panicle at onset of anthesis (third slide).

East Bijou Creek, Elbert County, Colorado. Mid-July; initiation of anthesis.

119. Unfulfilled florets- Spikelets of prairie sandreed showing florets at pre-anthesis stage. There is only one floret per spikelet in this species and the upper glume is considerably longer than the lower (Shaw, 2008, p.90). Both of these features were presented in these two photographs.

East Bijou Creek, Elbert County, Colorado. Mid-July; initiation of anthesis.

A Taste of True Prairie

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

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

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

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

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

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

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

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

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

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

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

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

Postclimax Tallgrass Prairie in the Southern Great Plains

To the west of the tallgrass prairie and true prairie regions of the humid and subhumid zones scattered "islands" of actual tallgrass prairie developed on sandy soils and, especially, sand dunes in the sermiarid zone. An edaphic complex of interacting phenomena including deeper and coarse-grained soils, greater usable water capacity of these soils, increased nutrient cycling and organic matter content, and more mesic (perhaps more fertile) microsites due to microrelief of dunes combine to permit development of potential natural vegetation that is more typical of range plant communities that form on more favorable (more moderate or less harsh or extreme) environments.

In the Clementsian monoclimax (climatic climax; zonal or regional climax) model such natural plant communities that develop on habitats more favorable than that characteristic of a vast spatial region over geological temporal scale are regarded and classified as postclimax. In the Tansian polyclimax and the Whittaker climax pattern models, communities on extra favorable habitats are regarded as just another of several climaxes due to edaphic and/or topographic features more than those of the prevailing climate. The end result is the same. The different sematics (including dogma) and underlying conceptual views depend on whether the potential terminal plant community (successional endpoint) is interpreted over the time frame of theoretical geologic cycles and a predicted, ultimate peneplane (monoclimax theory) or in the shorter temporal scale of soil formation on existing land forms or topography (polyclimax or climax pattern theory).

In the descriptions and discussion that followed in this section the designation of postclimax was used in preference to (instead of) edaphic, topographic, or edapho-topographic climax. The rational, logical basis for this choice was direct and simple. First, the doctrine of priority (first in time; first in rights or first use equals first claim) awards the title to postclimax not some polyclimax-based designation such as edaphic climax. Second, postclimax was the term used in the earliest descriptions of tallgrass vegetation on dunes in the Great Plains which was by University of Nebraska ecologists under leadership of Charles E. Bessey (Roscoe Pound, Frederic E. Clements, and John E. Weaver were the pivotal founders of Plant Ecology). Out of the Nebraska school (through Clements and Weaver and, later, Weaver students) emerged monoclimax theory. The first theoretical explanations of grassland vegetation development on extra favorable habitats like dunes or wetlands was that of monoclimax theory (ie. postclimax) as postulated and published by Clements.

Some ecologists, including range scientists, included dune grasslands on which the dominants were tallgrass species as mixed prairie (vs. tallgrass prairie) or, at least, as part of the larger, surrounding mixed prairie. Like the situation of monoclimax vs. polyclimax or climax pattern theory, the end result is the same whether the perspective is 1) dune grassland as a form, expression, or range type of mixed prairie or 2) tallgrass prairie on dunes or deep sand within the mixed prairie region. This latter view was incorporated in the following treatment with due acknowledgement of the designation, mixed prairie, by some workers and in description of the rangeland cover type (Shiflet, 1994).

The postclimax tallgrass prairie has an obvious ecological aaffinity with the proximate--frequently adjoining--sand sagebrush (Artemisia filifolia)-tallgrasss or midgrass savanna and the sand shinnery oak (Quercus havardii) scrubland. This affinity is most clearly seen in shared key decreaser grasses most notably sand bluestem (Andropogon hallii= A. gerardii var. hallii). Sand sagebrush is a widespread shrub throughout much of the semiarid Great Plains vegetation. This woody composite is a conspicuous member of climax range plant communities, including postclimax tallgrass prairies on dunes and deep sands. Relative species composition (cover, density, biomass, general abundance) of sand sagebrush varies from trace to modest proportions so that there is a continuum from grassland with widely scattered shrubs to a grass-shrub savanna to a sagebrush scrubland with something of an herbaceous (grassy) understorey. These sand sagebrush shrublands are usually to almost always abnormal woody invasions indicating range retrogression (for causes ranging from inadequate frequency of fire through overgrazing to previous farming followed by field abandonment). Brush invasion on degraded ranges notwithstanding, presence of shrubs like sand sagebrush, various scrub cherries (Prunus spp.), plains pricklypear (Opuntia polycantha) and subshrubs such as broom snakeweed (Gutierrezia sarothrae) are natural components of climax vegetation-- at small propotions (percentages) of total plant cover and biomass.

121. Postclimax tallgrass prairie- Mixed prairie on deep sand in the Southern High Plains consisting of two layers: 1) a tallgrass overstorey dominated by sand bluestem (Andropogon hallii) with little bluestem and upland switchgrass as associate species and 2) a mid-grass understorey with needle-and-thread and sideoats grama as dominants along with traces of sand dropseed and blue grama. Blue grama is usually thought of (and was treated elsewhere herein as) a shortgrass species, but on this Deep Sand range site it attained a height approaching that of sideoats grama and it was so scarce that it did not comprise a shortgrass layer in this grassland vegetation. Sand sagebrush (Artemesia filifolia) also occurred in trace proportions such that it did not add a shrub layer. This vegetation was thus grassland and not a mixed grass-sagebrush shrub savanna (a form presented immediately below). Forbs were not abundant, but wild alfalfa (Psoralea tenuifolia) was present.

Scale of this tallgrass form of mixed prairie that developed in a 17 inch precipitation belt can be guaged by the four-foot barbed wire fence. Shoots of sand bluestem remaining (and broken) from the previous growing season extend above the top wire. The short shoot or "culmless" sand bluestem had just started to elongate it shoots and was still in the pre-boot stage during the current growing season.

The rolling sand dune microrelief of this range site was shown conspicuously by featuring a dune in extreme left midground. In the photograph the dune sloped down from left (top of dune) to right (bottom of dune) so that the base level of the dune was directly in front of the second steel post from the left side of the slide.

Mixed prairie postclimax grassland dominated by tallgrass species, especially sand bluestem, was discussed in Weaver and Albertson (1956, ps. 261-265, 295-299).

The concept of postclimax will be confusing to both beginning students and "climax" students who were not instructed or self-taught in the Clementsian paradigm. It is helpful-- if not essential-- for students to have some grasp of the concept of postclimax (and other aspects of the Clementsian model such as preclimax and disclimax) because so much of the seminal and classical work on range vegetation, especially that of grasslands, was interpreted and described in the language of the Clementsian lexicon. The sand bluestem-dominated mixed prairie range presented here was but one example.

Aside from the need to be fluent in the prevailing language in which most of North American vegetation was described, concepts like postclimax remain useful in providing a rational explanation for development and persistence of many North American range types. Postclimax tallgrass prairie and mixed prairie dominated by tallgrass species (like sand bluestem) is one of the textbook examples of this. At regional scale the Sandhills Region of Nebraska and Colorado is one of the largest remaining units of postclimax vegetation while smaller areas of sandhills prairie like that featured in this slide are outliers of postclimax plant communities. Readers were referred to Weaver and Clements (1938, ps. 82-83, 85-86, 102, 110, 462, 475, 520-521).

Clements (1936, p. 269) used the sandhills tallgrass as an example of postclimax, but in a pattern characteristic of Clement's writing he did not offer a concise definition of postclimax. Allaby (1998) did: "In the monoclimax model of climax vegetation development, communities differing from the climatic climax, owing to cooler and/or moister conditions than are characteristic of the regional climate". For example, soils of the Sandhills permit faster infiltration rates and greater absorption of precipitation to deeper depths in the soil profiles which allows development of vegetation more mesic than that which is typical for the general region of the High Plains. Grassland vegetation of humid and subhumid regions can develop in a semiarid region due to more mesic soil conditions, a function of coarser-textured soils.

Southern High Plains. Cimarron County, Oklahoma. July. FRES No. 38 (Plains Grasslands Ecosystem). K-62 (Bluestem-Grama Prairie). SRM 721 (Sand Bluestem-Little Bluestem, Plains). Bluestem "Tall-Grass" Series, 142.11 of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Deep Sand range site. High Plains- Rolling Sand Plains Ecoregion, 25b (Woods et al., 2005).

Sandhills postclimax tallgrass prairie on eastern High Plains- Shown and described in the following section was an example of sandhills tallgrass prairie at the eastern margin of the Southern High Plains. This example was on a low dune form of this range cover type on the Agricultural Research Service Southern Plains Experimental Range in Harper County, Oklahoma. The range vegetation described below differed from that above in being more of the sand dune shrub-grassland in contrast to grassland on sandy soil of nearly level land. This was the difference between sand sagebrush-tallgrass prairie described below versus tallgrass-midgrass prairie presented immediately above. In Kuchler units these two climax range plant communities were Bluestem-Grama Prairie (K-62) shown above and Sandsage-Bluestem Prairie (K-63) shown below. The example of K-62 was on the High Plains section of the Great Plains physiographic province whereas the example of K-63 was on the Plains Border section of the Great Plains (Fenneman, 1931, fold-up map), a section later described variously as the Rolling Red Hills, Rolling Redlands, Rolling Red Plains (Frye and Swineford, 1949; Schoewe, 1949, ps. 302-307).

The Society for Range Management rangeland cover type that best fit this range vegetation on basis of location (SRM 722; written by a Southern Plains Experimental Range scientist) described this as sand sagebrush-mixed prairie. R.Rosiere adhered to traditional (=Clementsian) views and interpreted this as this as postclimax tallgrass prairie. Alternatively--but with the same net conclusion--this general range plant community could be interpreted as an edaphic or edapho-topogrphic climax if viewed from the polyclimax theory of Arthur Tansley or climax pattern interpretation of Robert Whittaker. (See Whittaker [1951, 1953] for detailed critique of these various theoretical views of climax plant communities.)

SRM 722 was written broadly enough to include general sand hills rangeland north through northeastern Colorado and southwestern Nebraska. Obviously these latter sand hills forms or variants of SRM 722 (and not to be confused with Nebraska Sandhills tallgrass paririe, SRM 602) that are dominated by species like needle-and-thread (Stipa comata) are mixed prairie in the literal (the textbook) sense. Sand hills and sand dune forms or variants dominated by sand bluestem and switchgrass are another major range type altogether with the difference being nothing less than that of tallgrass prairie versus mixed prairie. From the current author's perspective SRM 722 could--and probably should--have been split to draw this distinction. At least it could be specified that there is a tTallgrass prairie vriant versus a mixed prairie variant of Sand Sagebrush-Prairie (ie. leave Mixed out of the title). Other than for specified areas or subregions (and some species differences), SRM 720 (Sand Bluestem-Little Bluestem, Dunes) was a more apt designation than SRM 722 (Sand Sagebrush-Mixed Prairie) for sand bluestem-dominated grasslands on duneland in the Rolling Red Hills. Relative cover of sand sagebrush--and in climax in contrast to deteriorated-- range plant communities was another critical issue as shown clearly in beginning the title of SRM 722 with " Sand Sagebrush" .

This vegetation was obviously in the mixed prairie association, but it was an island of tallgrass prairie therein. The flora included major species of short grasses (buffalograss, blue grama) and mid grasses (sand dropseed, sand lovegrass, western wheatgrass, sideoats grama), but the species composition and community structure were those of tallgrass prairie. This was especially true for most of the interdunal habitats which were generally dominated by sand bluestem, the upland form of switchgrass, and little bluestem. Vegetation of dunes was often a microsite shrubland of sand sagebrush and Chickasaw plum (Prunus angustifolia) and perhaps some plants of Oklahoma plum (P. gracilis).

Some of the less mesic interdunal habitats supported such cover of sand sagebrush as to be a savanna, generally with herbaceous layers comprising mixed prairie. The range vegetation presented here, however, was obviously (at least to this rangeman) as a tallgrass prairie form or phase of sandhills grassland. As such it was included in the chapter deveoted to interior tallgrass prairie.

122. Tallgrass prairie on the Southern Great Plains- General views of postclimax tallgrass prairie in the eastern Great Plains. Physiogonomy, structure, and growth or life form (tallgrass of sod-forming and/or cespitose habit, both monocotyledon and docotyledon shrub) of range vegetation and gently rolling to undulating relief (topography) of plains land form were presented. Sand bluestem and little bluestem were the most common grasses, but there was also silver bluestem, Canada wildrye, sideoats grama, tumblegrass, western wheatgrass at local scale, and even patches of blue grama and buffalograss. Forbs were uncommon, but two species with a conspicuous presence were spectacle-pod (Dithyrea wislizenii var. palmeri) and thickleaf or aridland goosefoot (Chenopodium pratericola). Illinois bundleflower was also present but at lower density, cover, etc. than spectacle-pod and thickleaf goosefoot. Soapweed yucca (Yucca glauca) and sand sagebrush were present though infrequent on this pristine example of postclimax or edapho-topographic climax grassland.

More details of species composition and general aspects of the range environment followed in subsequent photographs and captions.

Southern Plains Experimental Range, Harper County, Oklahoma. Late June; late vernal to early estival aspect. FRES No. 39 (Prairie Ecosystem). K-63 (Sandsage-Bluestem Prairie). SRM 722 (Sand Sagebrush-Mixed Prairie) and/or SRM 605 (Sandsage Prairie): two competing rangeland cover types or two slightly different titles and descriptions of the same or very similar range vegetation. Bluestem "Tall-Grass" Series, 142.11 of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Central Great Plains- Rolling Red Hills Ecoregion, 27q (Woods et al., 2005).

123. More views- Range landscape and vegetation of postclimax tallgrass prairie on the Rolling Red Plains. Sandy soil and sand dunes provided an environment for grasses and shrubs that was more favorable (moister soil, more moderate microclimate) than that of range habitats which were more typical of this eastern margin of the Great Plains (the Plains Border section). More prominent relief (by Great Plains standards) and deeper rooting medium made for a more mesic and generally milder range environment which in turn resulted in development of a range plant community comprised of grasses having larger mature size (eg. tallgrass species such as sand bluestem, little bluestem, and switchgrass) and higher proportions of woody plants (greater crown cover of shrubs like sand sagebrush and Chickasaw plum). Hence, this has traditionally been interpreted as postclimax vegetation.

The first of these two photographs presented postclimax vegetation in an interdunal space, a fairly level land surface situated among sand dunes. Range vegetation in the local interdunal area was a dense stand of warm-season tallgrasses (switchgrass, sand bluestem, and little bluestem were all prominent) along with Canada wildrye and Illinois bundleflower. The two most conspicuous and generally most abundant forbs were spectacle-pod and thickleaf goosefoot. The second photograph showed another interdunal area (foreground) leading with an ever-so-gently elevational rise to a sand dune (background). In this second slide sand sagebrush joined the lineup of grass species listed for the first slide. Details of range vegetation on dunes followed in subsequent slides.

124. Studying range vegetation on the way to and at base of a sand dune- A synopsis view of postclimax tallgrass paraire on sandy land in the Plains Border section at eastern edge of the Great Plains was presented in the first of these two photographs. Sand bluestem, a decreaer tallgrass, was the overall dominant of the range presented here. Little bluestem and switchgrass were typically associate species except for local environments where they were dominant, even exclusive, species. Other grasses included Canada wildrye, silver bluestem, sand dropseed, sideoats grama, tumblegrass, blue grama, and buffalograss. The grass stand in foreground to midground of this first slide was so dense and with such heavy foliar cover of grass that there were almost no forbs present except for Illinois bunmdleflower. Shrubs were almost totally absent in this dense grass stand.

The second "photo-quadrant" included an east slope of a big circumference sand dune and a large interdunal area to the immediate right and rear of the featured slope of the dune. Even here on a hummocky low dune on edge of the High Plains, an east slope (even if just a slight inclination) was more mesic than adjacent microhabitats (environments formed by microrelief). The small patch of woody vegetation was mostly Chickasaw plum with sand sagebrush, the more xeric shrub, the associate species. At toe slope of the dune range vegetation (foreground) there was a local combination of Chickasaw plum and herbaceous plants with shoots of sand bluestem prominent. Other herbaceous species included the naturalized annual brome, Japanese chess, and western ragweed. The interdunal space was co-dominated by sand bluestem and the upland form of switchgrass.

There was a tremendous crop of biomass on this range, especially on the interdunal areas. This much standing crop was produced in a location receiving about 20 inches of average annual precipitation.

125. Studying range vegetation on a sand dune- Local stand of postclimax scrub on a sand dune in the Rolling Red Plains. This floristic cast featured woody dominants supported by herbaceous understudies. The first of these two slides showed most of a sand dune (the side and summit) that was typical for duneland range in this area. Sand sagebrush dominated sides of the dune while tallgrasses maintained control of soil at foot of the dune and Chickasaw plum (possibly some Oklahoma or slender plum) slightly edged out sagebrush to be "king of the hill" (or, rather, "of the dune").

Note on Prunus species: according to Atlas of the Flora of the Great Plains (McGregor et al., 1977) it was possible that there was both Chickasaw plum (Prunus angustifolia) and Oklahoma plum (P. gracilis) on sand dunes at Southern Plains Experimental Range.

The second slide featured the summit (if such term would apply) of the same dune. On this upper part of the dune plum was slowly pushing sand sagebrush downhill. Sand bluestem was the most abundnt grass and could be interpreted as an herbaceous associate species. Interesting biodiversity, including species and life or growth forms. Both woody and herbaceous plants were climax dominants on the range habitat of dunes, at least at roughly the compositions and locations shoen here. This would make for an interesting--though simple--gradient analysis.

126. Savanna form of postclimax sand sagebrush-tallgrass vegetation- Two views of Great Plains deep sand and dunes where warm-season tallgrassess (sand bluestem, little bluestem, and upland switchgrass) and miscellaneous other grass species (Canada wildrye, sand dropseed, silver bluestem, sideoats grama, blue grama, buffalograss, and Japanese chess) with sand sagebrush as the woody (shrub) component formed a savanna. This range plant community was postclimax when interpreted from monoclimax theory and/or an edaphic or edapho-topographic climax when viewed from polyclimax theory. Forbs were neither common nor indicator species. The two main (if uncommon) forbs were spectaclepod and thickleaf goosefoot. Illinois bundleflower ran a distant third.

In this range type and in this area in the Plains Border section at far eastrn edge of Great Plains there exist a continuum with regard to cover and density of sand sagebrush extending from a savanna with the dominant tallgrasses and grass species to a sand sagebrush shrubland with very little to almost no herbaceous understorey. The latter vegetation almost always indictes a highly degraded range that is often viewed as a disturbance climax in which range improvements (brush control and, often. range reseeding) are required to restore such depleted ranges.

The savannah form of range plant communities on Great Plains dunes and deep sand is often the potential natural vegetation (ie. postclimax range vegetation). That form or variant of duneland range vegetation with widely scattered plants (often local, fairly dense yet widely spaced stands) of sand sagebrush with even greater density and cover of decreaser tallgrasses was shown in these two photographs. Students will note that the sand bluestem was growing inside local stands of sand sagebrush where this dominant species stood taller than its climax shrub associate. It was underscored that sand sagebrush was climax at the approximate proportions (density, cover, biomass) of the range vegetation in these "photo-plots". A range plant community consisting primarily of sand sagebrush--a sagebrush scrubland or shrubland--constitutes a disclimax, especially when not "accompanied" by tallgrasses like sand bluestem, little bluestem, and switchgrass.

127. Sand country herbaceous cast- Composite views of major grass species on postclimax tallgrass prairie on semiarid duneland. Range vegetation featured here was in an interdunal space. Sand bluestem was the main grass species that made up most of plant cover in first photograph and was featured at shorter focl length in second photograph. Large shoots (including some of last year's mature-sized ones in first slide) and the unique light gray-green color set off sand bluestem from all other grasses. Some of the other grass species included an upland form (ecotype) of switchgrass, nodding or Canada wildrye, little bluestem, sand dropseed, buffalograss, and blue grama. Not all of these grasses were discernable in these two slides, but structure of this range vegetation and close-distance physiogonomy of it were presented, as they were representative of this postclimax tallgrass prairie.

An argument could be made that the species composition and architecture/structure of this range vegetation was a form of mixed prairie given the combination of tallgrass species like sand bluestem and switchgrass with shortgrasses such as blue grama and buffalograss. There were at least two glaring facts that weakened such an argument: 1) tallgrass species overwhelmingly dominated the vegetation and made up a disproportionately large percentage of the species composition based on biomass, foliar cover, physiogonomy (including aspect dominance), and even shoot density and 2) midgrasses like sideoats grama and western wheatgrass that are typically major charactaeristic components of mixed prairie were the least abundant group of grass species present on this range.

Note on orgainzation: Mixed prairie range (eg. climax vegetation dominated by needle-and-thread, blue grama, sideoats grama) in the sand hill area of the Central Great Plains were included in the Mixed Prairie chapter (also under the Grassland heading) of Range Types.

128. Basal shoots of sand bluestem (Andropogon hallii)- Examples of lower shoots (just above rootcrown) of sand bluestem growing on sandhills postclimax (= edaphic climax) tallgrass prairie in the Southern High Plains. Sand bluestem was the overall (and the defining) dominant tallgrass species of this climax prairie community. Little bluestem and switchgrass (upland form) were the associate tallgrasses with either of these species sometimes being the local dominant. These examples presented typical early season coloration of sand bluestemt along with close-up views of leaf blade and sheath. Note prominent parallel leaf veination.

Southern Plains Experimental Range, Harper County, Oklahoma. Late June; mid pre-bloom phenological stage.

129. A little higher up- Details of leaves and clum of sand bluestem growing on a sandhills postclimax (= edaphic climax) tallgrass prairie in eastern margin of the Southern Great Plains. Details of leaf collar and sheath were emphasized in this slide.

Southern Plains Experimental Range, Harper County, Oklahoma. Late June; mid pre-bloom phenological stage.

130. Goosed in the dunes- Thickleaf or aridland goosefoot (Chenopodium pratericola) growing in an interdunal area in postclimax tallgrass prairie in the southern sandhills of the Southern Great Plains. This specimen was growing on a range that could be described as a sand sagebrush-tallgrass savanna. Part of this postclimax range vegetation was dominated by sand bluestem and upland switchgrass whereas other local areas consisted mostly of mid- and shortgrass species or, on tops of sand dunes, scrub of sand sagebrush and wild plum. Thickleaf goosefoot is one of several Chenopodium species in the Great Plains Region, almost all of which (including this species) are annuals and of moderate palatability to range animals including grass -feeders like cattle. C. pratericola was a locally dominant forb in this deep sand vegetation.

Southern Plains Experimental Range. Harper County, Oklahoma. Late June.

131. Spectacle-pod (Dithyrea wislizeni= Biscutella wislizeni)- Spectacle pod is not only one of the most apt-named range plants it is also one of most common species of the Cruciferae native to the ranges of the Southern Great Plains and Tran-Pecos Basin and Range. Coulter (1891-1894) wrote of this species as native to the Staked Plains (High Plains portion of the Southern Great Plains) and sandy soils along the Rio Grande. Wooton and Stanley (1915) presented the species range as from Colorado and Utan to west Texas and south into the Mexican states. Spectacle-pod is thus a species of both Mixed Prarire and Shortgrass Plains grasslands as well as of the Chihuhuan Desert,It is especially abundant in the shinnery oak (Quercus harvardii) shrubland ecosystem and, at somewhat lesser cover and density, in the postclimax tallgrass prairie of the Southern Great Plains. Spectacle-pod has a high preference for habitat with sandy soils.

Spectacle-pod appears to have a very versatile life cycle having been described by various taxonomists as annual, biennial, or perennial. This pattern of resource allocation is characteristic of plants of harsh habitats. Whle conducting reserch for his Master's thesis this author found spectacle-pod to one of the most densly populated of the larger forb species on the disclimax Chihuhuan Desert of the New Mexico State Universtiy College Ranch. This was during a period (the years of 1972-1973) of average or higher precipitation. Sebsequent visits to the same study ranges of the College Ranch following dry periods and severe drought revealed little and no evidence, respectively, of this cricifer. These findings were consistent with an opportunistic range species.

The specimen shown here was growing in a typically dense population in a sand shinnery oak (Quercus haavardii) shrubland in which there had been a recent surface fire. (Note small shinnery oak resprouts.)

Beckham County, Oklahoma. June.

132. Inflorescence and fruit of spectacle-pod- The fruit type of is a silique, defined by Smith (1977, p. 307) as a broad, longitudinally dihiscent fruit in which fruit walls part from a persistent internal partition called the replum.

The photographs of the flower cluster showed the prominent and clearly separate or unjoined four petals of the corolla.

Beckham County, Oklahoma. June.

133. Edge of an island of tallgrass prairie in a High Plains "sea" of mid- and shortgrasses (with some scrub)- Two photo-plots of postclimax tallgrass (sand bluestem and little bluestem) prairie at the perimeter of a savanna of sand shinnery oak with sand bluestem and little bluestem that developed as an ecotone between tallgrass prairie and sand shinnery oak range ecosystems. At this point the savannah was to the rear with advance into the tallgrass prairie at peak standing crop and grain-ripe with onset of dormancy in bluestems.

This first slide featured the surface of the Tivoli deep sands soil. This slide was taken under a cloudless sky with full sun. The second of these photographs showed sand bluestem and little bluestem on the shallower Brownfield soil. This second photograph was taken under an overcast sky and showed the pale pubescence of bluestem inflorescences to advantage (but with less depth of field than in the full-sun shot). Here at the edge of grassland and away from savannah and, farther yet, shrubland the only shrub of much consequence was broom snakeweed, a local associate species. Another shrub, though one of much less cover and density than broom snakeweed, was sand sagebrush. Sagebrush was also present in the sand shinnery oak shrubland as seen above. Southwestern (Chyrsothamnus pulchellus) was a widely scattered shrub present in trace amounts in both the savanna and grassland and not, to any notable extent, in the shinnery oak range ecosystem.

Organization note: Viewers wishing to see changes in range vegetation from sand shinnery oak scrub duneland through the the ecotonal savanna to this tallgrass prairie can find the relevant photographs under Shrublands (Shinnery Oak).

Weaver Ranch, Roosevelt County, New Mexico. Autumnal aspect, October. Early dormancy and fruit-ripe pehnology of climax grasses. In theory this grassland would be (or could be seen as) FRES No.39 (Prairie Grassland Ecosystem) and K-66 (Bluestem Prairie) as a postclimax or a smaller-scale edaphic climax in a region the zonal vegetation of which is FRES No. 38 (Plains Grassland). This is problematic however because there is no Kuchler unit of tallgrass prairie in FRES No. 38. Itstead this was mapped as K-58 (Grama-Buffalograss). SRM 720 (Sand Bluestem-Little Bluestem [Dunes]). Bluestem "Tall-Grass" Series 142.11 in Plains Grassland biotic community 142.1 of Brown et al. (1998). High Plains- Llano Estacado Ecoregion, 25i (Griffith et al., 2006).

134. Postclimax prairie- Tallgrass prairie of sand bluestem and little bluestem in a semiarid zone in which the region climax range vegetation is mixed prairie and where there is far more "shortgrass country" than tallgrass range. This pair of photographs provided another study in light and color contrasts resulting from different conditions of natural light. The first slide (taken in full sun; under a completely cloudness sky) was of a stand of postclimax tallgrass prairie that was a consociation of sand bluestem, for all practical purposes the only grass species present in this stand. The shrub was broom snakeweed and the forb (a specimen in right, immediate foreground) was annual wild buckwheat (Eriogonum annuum). The second slide (under an overcast sky) featured a range community of more species diversity including sand bluestem, little bluestem, Indiangrass (Sorgastrum nutans), broom snakeweed, and annual wild buckwheat. In the instance of this second slide, grass panicles with their pale or silvery pubescence showed up to much better advantage under conditions of reduced light.

Range photographers frequently benefit from different light conditions depending on what aspects and characteristics they wish to feature. Atmospheric conditions with cloud cover that varies dramatically over a short period of time, while a source of aggravation if not frustration (especially with moment-to-moment light dynamics) to the shutterbug, is ideal for most kinds of range photography. These two photographs and the two in the preceding set provided examples of that lesson.

Anyway, this was a beautiful postclimax tallgrass prairie amid an otherwise "vast sea" of mixed prairie. Even the crotchiest ole range professor can fight at least half a semester's battle with university Pharisees treasuring scenes like these in his soul.

Weaver Ranch, Roosevelt County, New Mexico. Autumnal aspect, October. Tallgrass species were at pehnological stage of grain-ripe and beginning dormancy. In theory this grassland would be (or could be seen as) FRES No.39 (Prairie Grassland Ecosystem) and K-66 (Bluestem Prairie) as a postclimax or a smaller-scale edaphic climax in a region the zonal vegetation of which is FRES No. 38 (Plains Grassland). This is problematic however because there is no Kuchler unit of tallgrass prairie in FRES No. 38. Itstead this was mapped as K-58 (Grama-Buffalograss). SRM 720 (Sand Bluestem-Little Bluestem [Dunes]). Bluestem "Tall-Grass" Series 142.11 in Plains Grassland biotic community 142.1 of Brown et al. (1998). High Plains- Llano Estacado Ecoregion, 25i (Griffith et al., 2006).

135. In prairie and in scrub- Sand bluestem is the climax dominant grass (major decreaser) species on both postclimax tallgrass prairie (first or upper slide) and sand shinnery oak scrubland (second or lower slide). Shoots of sand bluestem in these (and above) photographs were in dormancy or near-dormancy stage with ripe grain. The associate species was the subshrub, broom snakeweed.

Weaver Ranch, Roosevelt County, New Mexico. Autumnal aspect, October. Early dormancy; grain-ripe phenology.

136. Details of the dominant decreaser grass- Sand bluestem was the dominant herbaceous species on sand shinnery oak duneland range (Tivoli deep sands) and co-dominant with little bluestem on tallgrass prairie that developed near shinnery oak shrubland.

Weaver Ranch, Roosevelt County, New Mexico. Autumnal aspect, October.

137. Sand bluestem-little bluestem-sand sagebrush- A sand dune cover type. Clements and Weaver (1938) regarded such communities as postclimax, a tallgrass variant type in the mixed prairie region. This cover type is shown in the grassland section because it is a grassland type and is not a sand sagebrush-sand bluestem savanna. This type occurs exclusively on deep sand sites and is interspersed among blue grama-buffalograss, western wheatgrass, and other forms of mixed prairie and shortgrass plains grasslands.

Chaves County, New Mexico. June. FRES No. 39 (Prairie Ecosystem). K-63 (Sand Sagebrush-Bluestem Prairie). SRM 720 (Sand Bluestem-Little Bluestem, Dunes).

Another Taste of Tallgrass Prairie in the Mixed Prairie Region

Developing "smack-dab" in the heart of mixed prairie there was, in Clementsian monoclimax thoery, a postclimax tallgrass prairie. In polyclimax theory of Tansley and climax pattern theory of Whittaker, this range vegetation would be interperted as an edaphic climax. This grassland that was more mesic than that of surrounding mixed prairie had developed on deep, sandy soil in what is known as the Great Bend Sand Prairie. Regardless of theoretical perspective, this tallgrass sand prairie was dominated by sand bluestem (Andropogon hallii) and with Indiangrass and switchgrass as associates. An example of this sand bluestem-Indiangrass-switchgrass prairie that was Excellent condition class was growing "just down the road" from the mixed prairie range stocked with heavy steers presented immediately above.

This postclimax, edaphic tallgrass prairie was included at this juncture to illustrate to students how vastly different range sites can be over remarkablly small distances and how this can result in different range cover types developing adjacent to each other. This example was also included in the postclimax tallgrass prairie portion of Mixed Prairie-II herein.

138. Bigger and taller grass on deeper sand- Postclimax tallgrass prairie dominated by sand bluestem with switchgrass, Indiangrass, sideoats grama as local associate species. Other abundant grasses included little bluestem and blue grama. The dominant forb in this early summer society was fineleaf or cutleaf hymenopappus (Hymenopappus filifolius). This range vegetation was essentially edaphically determined with the relatively deep sandy soil being conducive to a more mesic range plant community than that of adjoining mixed prairie that was presented in previous (above) photographs.

Kiowa County, Kansas. Late June (early estival aspect). FRES No.39 (Prairie Grassland Ecosystem) and K-66 (Bluestem Prairie) as a postclimax or a smaller-scale edaphic climax in a region the zonal vegetation of which is FRES No. 38 (Plains Grassland). This is problematic however because there is no Kuchler unit of tallgrass prairie in FRES No. 38. Itstead this was mapped as K-62 (Bluestem-Grama Prairie). SRM 721 (Sand Bluestem-Little Bluestem [Plains]). Bluestem "Tall-Grass" Series 142.11 in Plains Grassland biotic community 142.1 of Brown et al. (1998). Sands range site. Central Great Plains- Great Bend Sand Prairie Ecoregion, 27c (Chapman et al., 2001).

139. Skinny plainsmen- Columbia slimleaf, fineleaf hymenopappus, or cutleaf hymenopappus (Hymenopappus filifolius) was the major forb on this post-climax tallgrass prairie tht developed on deeper, sandy soil on the eastern portion of the Southern High Plains. Kiowa County, Kansas. Late June, peak bloom.

[ Home ]