Meadows and Related Marshes

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Marshes are those wetlands dominated by herbaceous vegetation—primarily grasses, sedges, rushes, cat-tails, etc.—growing in soils more or less permanently waterlogged or at state of saturation (ie. soils with water above field capacity most of the time). Vegetation growing on land covered with water permanently not periodically. Vegetation is climax at climatic time-scale; it is the potential natural wetland vegetation that is in dynamic equilibrium with the determinative factor(s) of habitat be it climate, soils, fire, flood, etc.

Marshes have traditionally been divided into two or three categories based on salinity of water:

Freshwater Marsh,
Saltwater Marsh and, sometimes,
Brackish Marsh (for those with mixing of upstream fresh water with saline water from oceans, salt lakes, etc.

These categories or general kinds of marshes include several distinctive wetlands with such freshwater marshes as everglades, tule marshes, and borders of oxbow lakes and such saltwater marshes as smooth cordgrass all along the Gulf and Atlantic Coasts or tidal marshes in bays.

Meadows shown as range cover types are those that conform with definition (2) of the Society for Range Management (Kothmann, 1974; Jacoby, 1989):

"Openings in forests and grasslands of exceptional productivity in arid regions, usually resulting from high water of the soil, as in stream-side situations and areas having a perched water table."

These are of two major or fundamental types:

Wet Meadow- "A meadow where the surface remains wet or moist throughout the summer, usually characterized by sedges and rushes [less commonly by hydrophytic grass species]" and
Dry Meadow- "A meadow dominated by grasses [less commonly bysedges or other grasslike plants adapted to less mesic or hydric conditions] which is characterized by soils which become moderately dry by mid-summer" (Kothmann, 1974; Jacoby, 1989 as [amended by Rosiere herein]).

Meadows are defined as more or less natural (comprised of native species) kinds of vegetation (or biotic communities if animals included) or ecosystems irrespective of the use made of them. In other words, as used herein meadows are like grasslands, deserts, forests, tundra, etc. They are biomes (of admittedly small spatial scale) delimited by the predominant growth or life form of vegetation. Meadows, grasslands, savannas, etc. are not —as is range by contrast— a kind of land based on it’s use (eg. any vegetation is by definition range only when it is subject to grazing or browsing). Meadows are meadows whether or not they are grazed or mowed for hay. Meadows become range (rangeland cover types) when used for grazing (or mowing). SRM definition (1) of meadow specified a primary use "for hay production" (ie. hay meadows). Many meadows represented by the vegetation displayed herein do qualify under both definitions, but it is only definition (2) that is applicable as basis of natural vegetation. Otherwise tallgrass prairies used for production of prairie hay— which are hay meadows only — would be meadows. This would be misleading because hay meadows are meadows only by criteria of definition (1) (eg. bluestem prairies can only be hay meadows and they should not be confused with meadows based on soil water criteria).

Marshes are some of the most valuable of all wetlands. They are also some of the productive (both ecologically as in Net Primary Productivity and agriculturally as in biomass production for hay, natural pasture, and wildlife habitat). This is especially so for waterfowl (ducks, geese, cranes, herons, plovers, swans, etc.). They are also important breeding grounds for fisheries, including as oyster beds. Marshes include wet meadows (there is not always a clear distinction between wet and dry meadows as some are intermittent or ephemeral wet meadows, especially subalpine mountain meadows). Inland wet meadows on floodplains are one extremely important sources of hay for beef cattle in the Intermountain Region. With careful and conscientious management such native meadow or tule (= bulrush) hay can be produced on flood meadows serving as prime waterfowl habitat.

Note on arrangement: Coastal marshes such as those in the Gulf of Mexico and along the Pacific Coast were included and dealt with in chapters deveoted to grasslands and related herbaceous vegetation associated with those marshes rather than being included in this chapter. For example, the Coastal Prairies and Marshes vegetational zone of Texas and Louisiana form an integral and inseperable unit (at both ecosystem- and landscape-scale) in which fresh, saline, brackish marshes and various types of tallgrass prairie exist as mosaics of range plant communites. These wetland range cover types were not included in the present chapter.

In the same vein, playa lakes are some of the most valuable wetlands in North America. Some of these (at least some aquatic zones of them) could be regarded as marshes. These natural wetlands are, however, integral parts of the surrounding mixed (mid-grass) prairie or shortgrass plains, often at levels or degrees of integration that do not exist, say, between mountain meadows and surrounding forests. For that reason playa vegetation was included in chapters devoted to those grasslands and not in the current chapter.

1. Narrowleaf cattail (Typha angustifolia) and American bulrush (Scirpus americanus) marsh surrounded by squirrel bottlebrush and inland saltgrass, which in turn is surrounded by a zone comprised of sideoats grama and little bluestem. Historic Bents Fort to far right, Otero County, Colorado. July. FRES No. 41 (Wet Grasslands Ecosystem). K-42 (Tule Marsh) in middle of K-62 (Bluestem-Gramagrass Prairie). No SRM listing for this cover type (or for any prairie marsh type). A composite of Rush Series and Sedge Series in Plains Interior Marshland biotic community of Brown et al. (1998). Southwestern Tablelands- Piedmont Plains and Tablelands Ecoregion, 26e (Chapman et al., 2006).

2. The cat and her tail- Narrowleaf cattail in peak bloom showing the pistillate infloresceence, "cat" (below) and staminate inflorescence, "cat tail" (above).

Otero County, Colorado. Late June; anthesis.

3. "Nervous as a cat in a room full of rocking chairs"- Close-up view of the inflorescence of the narrowleaf cattail presented in the immediately preceding slide. Emphasis was on the "cat tail" portion, the male inflorescence (or male portion of the inflorescence) shown in its full length above a small part of the upper portion of the "cat", the female part of the flower cluster or the pistillate inflorescence (depending on interpretation). Typha species are interpreted as monoecious so that there are separate male and female inflorescences even though they adjoin each other on the same stalk. Smith (1977, p. 239) described this arrangement as "a tightly compacted terminal spadix". Now ain't that the cat's whiskers (or something else)?

Otero County, Colorado. Late June; anthesis.

4. Common or broad-leaf cattail (Typha latifolia)- This is one of three Typha species (as commonly interpreted by most authors) found on wetlands of the central grasslands of North America. Along with members of the rush and sedge family (and some minor ones like the burr-reed family) cattails constitute the groups of range plants commonly known as grass-like plants. This group or category of range plants carries no official taxonomic recognition in the lexicon of Plant Systematics, but is used primarily by rangemen and foresters to designate those species of monocots which do not produce flowers with showy or conspicuous petals and yet which are not grasses. These include several orders of monocotyledons. Vascular spore-bearing plants such as the horsetails or scouring rushes (Equisetum spp.) and monocots with obvious petals (eg. the orchids) are classified as forbs by natural resource practitioners like rangemen and wildlifers. foresters. The grasses and grass-like plants are, in turn, known collectively as graminoids.

Cattails are some of the most common and conspicuous species of grass-like plants on both marshes and wet microsites on grasslands across North America. Cattails grow not only in natural depressions like potholes and wet draws but also around man-made structures that impond water such as ditches, lakes, sewage lagoons, and farm and ranch ponds such as the one shown here on Kelly or Lone Oak Prairie, Craig County, Oklahoma. June.

5. Inflorescence of common cattail- Cattail gets its common name from its prominent flower cluster composed of distinctly separate groups of staminate and pistillate flowers. The upper structure (seen here as the gray, curved portion) of the inflorescence consist of male flowers and is called the "cattail"; the lower structure (the brown, cylindrical part) of the inflorescence is composed of female flowers known as the "cat".

6. Showy wetland forb- Showy milkweed (Asclepias speciosa) is a common botanical member of meadows throughout much of the Great Plains. It is one of the most widely distributed milkweeds in North America with a species range extending from the Great Plains (Texas panhandle to southern Saskewan ) and westward to the Pacific Ocean (Burrows and Tyrl, 2001, p. 129). The pretty specimen was captured in early morning light growing with narrowleaf cattail in a wetland in southwestern Colorado.

Asclepias species have traditionally been divided into two groups based on width of leaf: 1) broad-leaved and 2) narrow-leaved. Showy milkweed is one of the broad-leafed species. A. speciosa is one of the many Asclepias species that can be a stock-poisoning plant under the right (or wrong) conditions. Readers were referred to Kingsbury (1964, ps. 267-271) and Burrows and Tyrl (2001, ps. 125-135). Research showed that the toxicity of milkweeds is due (at least mostly) to compounds characterized as asclepiad cardiotoxins and generically called cardenolides. Toxicity of among specific milkweed species varies considerably with potential for animal poisoning assumed to be strongly related to cardenolide content (Burrows and Tyrl, 2001, ps. 131-133). A. speciosa is intermediate in concentration of cardenolide (Table 12.1 in Burrows and Tyrl, 2001, p. 133).

Otero County, Colorado. Late June; peak bloom stage.

7. Portion of a stand- Portion of a large population (a colony) of three square, American bulrush, or chairmaker's rush living in the shallow, channel of Rush Creek, an emphemeral stream in the High Plains.

Lincoln County, Colorado.Early July, fruit-ripening stages.

8. Three square group- Group or cluster of floral units with ripening fruit of three square, chairmaker's rush, or American bulrush. In species in the Cyperaceae individual flowers are grouped into units of organization called spikelets. Four spikelets of Scirpus americanus were visible on the triangular-shaped shoot.of this specimen. Each flower has only one glume and can produce only one fruit, the fruit type of which is an achene.

Rush Creek, Lincoln County, Colorado. Early July, fruit-ripening stages.

Continuum of Wet Prairie to Marsh

A mosaic of wetlands ranging from wet prairie to river floodplain to marsh exist in the immediate vicinity of the Salt Fork of the Arkansas River in northern Oklahoma and southern Kansas. This natural "patchwork" of various grasslands and marshes provided a good example herbaceous wetlands within the tallgrass prairie region. Marshes are not grasslands strictly speaking or in precise usage, but in the example provided here (and it was a typical situation over formerly large areas in the once-vast grassland domain of central North America) marshes and wet prairie formed an intricate, inter-related herbaceous vegetation at both landscape and ecosystem levels. These two biomes were interconnected by processes at landscape-scale (an example worthy of an undergraduate textbook in Landscape Ecology). For this reason these two general plant communities (grassland and marsh biomes) were treated simultaneously and included in this location in Range Types of North America. The tule marshes were also included under the Meadows chapter of the Grassland biome to facilitate use by students.

In the Arkansas River lowland example employed below differences in salinity and moisture conditions of soil at small (local) spatial scale resulted in a small-scale mosaic of wet (and somewhat saline) tallgrass prairie, mixed prairie, and bulrush or tule marsh in amazingly close proximity to each other.

Wet saline prairie and tule marsh had developed side-by-side. They constitute grassland and marsh and so were essentially of two different biomes. These two range plant communities were interconnected by processes at landscape-scale (ie. a textbook example of Landscape Ecology). The tallgrass prairie was treated under the Tallgrass (Interior-II) chapter of the Grasslands biome.

9. Wet Saline Prairie- Tallgrass prairie comprised of switchgrass, the dominant, and big blusestem, the associate species, surrounded and, in turn, was surrounded by more saline areas dominated by inland saltgrass (Distichlis stricta= D. spicata var. stricta) with associated species ranging from the forb, frogfruit (Lipppia lanceolata) which was the broadleaf species in foreground of this slide, to plains lovegrass (Eragrostis intermedia) to Japanese brome (Bromus japonicus) and, with much less cover, cheatgrass (B. tectorum). Both of these Bromus species are naturalized Eurasian cool-season annuals.

The stand (a consociation) of switchgrass (the bottomland form; a lowland eoctype) seen in background background was shown at local scale in the next photograph. The local stand of tallgrasses (mostly switchgrass) in left foreground was on a more saline microhabitat so that grass growth and development was less and behind that of the same species on less saline soil.

Clarification: It was explained in several captions below that grassland vegetation on the Arkansas River lowland that was described in this section was labeled as wet prairie or wet saline prairie and not marsh. Designation of marsh was applied only to wetlands having their land surface covered with water during most--at least during critical parts--of the plant-growing season, and typically supporting grasslike plants rather than grasses. The tule or bulrush marsh presented later provided an example of marsh vegetation. Such marshes are not tallgrass prairie (they are not grassland at all), but the example referred to was included (below) in this portion of the Tallgrass Prairie chapter because these herbaceous wetlands occur in restricted areas within the tallgrass prairie region.

General floodplain (lowland) of the Salt Fork of Arkansas River. Great Salt Plains National Wildlife Refuge, Grant County, Oklahoma. June; late vernal aspect. Range plant communities of tallgrass species (mostly big bluestem and switchgrass) was FRES No. 39 (Prairie Grassland Ecosystem). K-66 (Bluestem Prairie). SRM rangeland cover type 710 (Bluestem Prairie), but situated within this range community were communities of short- and/or midgrass species, especially inland saltgrass. These latter grassland communities often covered greater area than surrounding tallgrass vegetation. There was not an SRM inland saltgrass rangeland cover type. Bluestem "Tall-Grass" Series 142.11 of Plains Grassland biotic community, 142.1 (Brown et al., 1998, p. 40). Saline Subirrigated range site. Central Great Plains- Prairie Tableland Ecoregion, 27d (Woods et al., 2005).

10. Standing together- Stand of tallgrass prairie, of which a bottomland ecotype of switchgrass was dominant and big bluestem was associate to local co-dominant, on higher ground of a wet prairie (floodplain of Arkansas River). This was a "photo-quadrant" of the range vegetation shown in the background of the immediately preceding photograph. Tallgrass stands like this one had developed on the higher level (elevation) land whereas lower ying local relief commonly supported consociations of inland saltgrass. Certain plant species such as plains lovegrass, Illinois bundleflower, and both japanese chess and cheatgrass (two naturalized Eurasian annual grasses) were more common in local ecotones (edges) between with these two distinct range plant communities. This extensive vegetational mosaic existed around freshwater tule (bulrush) marshes that had stands of eastern cottonwood (Populus deltoides) around their perimeters. This spatial arrangement of freshwater marshes and a "patchwork" of tallgrass wet prairie interspersed with midgrasses on the greater floodplain of the Arkansas River rproduced a landscape-scale grassland-marsh complex with interacting ecosystems.

11. Tallgrass and midgrass; mound and swale- Two views of local stands of tallgrass wet prairie (switchgrass and big bluestem, with the former more commonly dominant) on slighted elevated ground (including mima mounds as shown here) interspersed with local, primarily single-species stands (consociations) of inland saltgrass on lower elevation land. Differences in elevation of land (soil surface) was usually just a matter of inches to a few feet even from depth of swales or "micro-valleys" to top of mima mounds, but it was sufficient to account for the profound differences in these local range plant communities. Salt content was undoubtedly a major factor in determining which of these two drastically distinct range plant communities grew on the two forms of local relief (micro-topography). Other edaphic factors were almost assuredly involved also, including greater soil depth on mjma mounds.

The first photograph presented a view of this range at a camera distance that showed the overall vegetational mosaic and "lay of the land" with both mima mound and low-lying intemound spaces. The second photograph was taken at shorter camera distance and showed specifics of the local relief and corresponding range vegetation (eg. switchgrass and big bluestem on the same mima mound and inland saltgrass all around base of teh mima mound). Relatively large areas of bare soil surface were widespread and characteristic of this wet prairie range.

Illinois bundleflower and plains lovegrass were locally abundant at outer edges of the stands of midgrass (mostly inland saltgrass) such as shown growing conspicuously in center and right foreground in the second slide. Western ragweeed (Ambrosia psilostachya) was an associate species on such perimeters (lower right corner of second slide).

Question as to proper designation of wetland: It was not known whether this natural wetland vegetation was more precisely described as wet prairie or as salt marsh. Perhaps the tallgrass (switchgrass and big bluestem) range vegetation was wet prairie and the slightly lower elevation and more saline soil (usually a consociation of inland saltgrass) was salt marsh. This author was reluctant to label any grassland plant community as a marsh instead restricting the designation of marsh only to land inundulated with water (standing water on the land surface) for a good part of the plant-growing season (eg. the tule or bulrush marsh covered below).

12. Salty stand- Wet saline prairie in part of the Salt Fork Arkansas River Valley that was a consociation of inland saltgrass that extended over a large area as part of a vegetational mosaic with tallgrass (switchgrass and big bluestem) prairie that developed on slightly land. A large expanse of this tallgrass plant community was in the far background of this photograph. About the only other plant species in this expansive stand of inland saltgrass was Japanese brome or Japanese chess, and it was present only as scattered individuals.

Successional state of the inland saltgrass community was not known, but this worker suspected that it was climax, or at least the potential natural vegetation for which a climax might not exist if the soil in the swale (the fairly level land situated within or among mima mounds) was not a mature soil. The description of this range site in the soil survey (Soil Conservtion Service, 1985, p. 55) stated that "continued overgrazing and extreme climatic conditions" ultimately results in range deterioration to a degraded community including inland saltgrass, ragweed, and annual grasses. The pattern o f retrogression on this range site (includinga brief description of intermediate successional stages) may well be an accurate visualization of the path of range depletion, but that range site description did not--indeed, by itself, could not--explain simultaneous presence of extensive populations (single-species stands) of inland saltgrass immediately adjacent to the obvious tallgrass climaax of switchgrass and big bluestem.

It appeared to this rangeman that both the tallgrass and the inland saltgrass communities were climax . Any overgrazing (it would have been in the somewhat distant past as this grassland was not being overgrazed and had not been overgrazed in recent years) would have resulted in replacement of the obvious tallgrass climax on mima mounds the same as on intermound spaces. Thus, it seemed highly unlikely that inland saltgrass was other than climax range vegetation (ie. a consociation). The key words in the range site description by agency range conservationists (Soil Conservtion Service, 1985, p. 55) were most likely "extreme climatic conditions". In more precise ecological terms these three words would be read as something like "harsh microclimate" (ie. "climatic conditions" would refer to microhabitat, microsite, or microenvironment) where much of the "extreme" nature of conditions would edaphic and/or topographic (perhaps reflecting drainage, salt accumulation, and related factors). Presence of climax (decreaser) Illinois bundleflower along with "weedy" seral forbs like western wheatgrass in stands of inland saltgrass was further evidence of the climax nature of range plant communities dominated by inland saltgrass. It was illogical to conclude that overgrazing had depleted tallgrasses and resulted in their replacement by inland saltgrass on land of level or flat microtopography while right next to this microland form tallgrass (switchgrass and big bluesetm-dominated) vegetation on mima mounds had not been impacted by grazing, or had recovered from past grazing abuse so much faster. Some other factor(s) had to be more responsible than grazing management.

General floodplain (lowland) of the Salt Fork of Arkansas River. Great Salt Plains National Wildlife Refuge, Grant County, Oklahoma. June; late vernal aspect. FRES No. 41 (Wet Grasslands Ecosystem). No Kuchler or SRM designation for inland saltgrass. In Brown et al. (1998, p. 45) this would be Saltgrass Series 242.34 (if one was shown which it was not) of Plains Interior Marshland 242.3. Saline Subirrigated range site. Central Great Plains- Prairie Tableland Ecoregion, 27d (Woods et al., 2005).

13. Fresh water bulrush or tule marsh- General or overall views of a freshwater marsh at edge of the Arkansas River Valley (Salt Fork) existing as a consociation of tule or bulrush known variously as Americn bulrush, saltmarsh bulrush, Olney threesquare, and chairmaker's club-rush, (Scirpus olneyi= S. americanus= Schoenoplectus americanus) and with the twining forb, fogrfruit (Lipppia lanceolata), as associate species. Minor (other than locally) plant species included American water plantain (Alisma subcordatum), southern annual saltmarsh aster (Aster divaricatus= A. subulatus var. ligulatus), and Engelman's spikerush (Eleocharis engelmannii). For all practical purposes there were essentially no other plant species present in this freshwater marsh.

The first of these two slides presented a wider view of the entire range plant community of the tule marsh and associated range vegetation around the perimeter from adjoining plant communities. Range communities around the margins of the marsh wer shown in the second slide. This second photograph showed a local forest of eastern cottonwood that had developed around edges of the marsh. This local forest had been invaded by the naturalized and dreadfully invasive shrub, Russian olive (Elaeagnus angustifolia), that was readily distinguished by its silvery gray-colored leaves.

Edge species: in this immediate locality there were places where sedges (Carex and Cyperus spp.) and rushes (Juncus spp.) grew along perimeters of freshwater tule marshes where this range vegetation contacted wet--often saline--prairie. An example of such contacts and the resulting local-scale ecotonal (transitional) vegetation, including sedges and rushes, was shown at end of this section.

Great Salt Plains National Wildlife Refuge, Grant County, Oklahoma. June; late vernal aspect. FRES No. 41 (Wet Grasslands Ecosystem). K-42-Tule Mrashes. No SRM designation of a rangeland cover type for tule marshes. In Brown et al. (1998, p. 45) this would be Bulrush Series 242.33 (if one was shown which it was not) of Plains Interior Marshland 242.3. Central Great Plains- Prairie Tableland Ecoregion, 27d (Woods et al., 2005).

14. Stand of chair-makers' rush ( Scirpus americanus)- The local freshwater marsh introduced in the preceding two photographs was a consociation of a major species of bulrush that has a "pasal" of both common and scientific names including Olney threesquare, chairmaker's club-rush, saltmarsh bulrush or Americn bulrush (Scirpus olneyi= S. americanus= Schoenoplectus americanus). Frogfruit was the associate--and about the only other--plant species of this marsh other than incidental (found only sporatically) species which were listed in the immediately preceding caption. It was possible that there were some infrequent plants of hardstem bulrush (Scirpus acutus= Schoenoplectus acutus), but this species could not be positively identified.

Note on taxonomy of Cyperaceae including the bulrushes or tules: Radical changes in nomenclature and general taxonomic organization have been in the Cyperaceae that have created confusion, if not chaos, among all users of scientific names except for the elites who change the names and systematics. To ease the pain and facilitate study of range vegetation both the traditional and revised (ie. revolutionary) binomials were shown for this species.

Detailed views of this wetland range vegetation were presented in the next two slides and caption.

Grant County, Oklahoma. June; late vernal aspect. FRES No. 41 (Wet Grasslands Ecosystem). K-42-Tule Mrashes. No SRM designation of a rangeland cover type for tule marshes. In Brown et al. (1998, p. 45) this would be Bulrush Series 242.33 (if one was shown which it was not) of Plains Interior Marshland 242.3. Central Great Plains- Prairie Tableland Ecoregion, 27d (Woods et al., 2005).

15. Those of a freshwater marsh- Two progressively closer views of range vegetation in a freshwater marsh almost exclusively dominated by chairmakers' club-rush, chair-makers' rush, Olney threesquare, or Americn bulrush. The associate species was frogfruit (mostly visible in the second slide). Other--though only incidental--species included American water plantain, southern annual saltmarsh aster, and Engelman's spikerush. The chairmakers' or American bulrush was in early bloom stage. (An example of inflorescence and stem of this bulrush species was presented below.)

16. Tule marsh and pond (and cows in it)- A vegetational mosaic of tallgrass (switchgrass and big bluestem-dominated) wet prairie, inland saltgrass saline prairie, and freshwater marsh dominated by (a consociation of) American bulrush or chairmakers' rush vegetation on floodplain (lowland) of the Salt Fork of Arkansas River. These three distinct range plant communities were shown and described above (in this section, entitled Continuum of Wet Prairie to Marsh). The tallgrass (switchgrass-big bluestem) wet prairie and the inland saltgrass low (saline) prairie existed in such intiricate and intimate association that they could be viewed as one general grassland (a biome) community that was distinct from the marsh (another, even if small-scale, biome). That these two (if viewed at biome-scale and distinction) or three (if viewed on basis of dominant plant species) range plant communities sometimes developed "cheek by jowl" was shown cleaerly in this photograph. In fact, a natural (not manmade) freshwater pond (with two cows) in the tule marsh was thrown in for good measure.

Range vegetation in foreground was transitional (ecotonal) between tallgrass wet prairie and tule (bulrush) marsh. Major range species in this local ecotone included switchgrass (a bottomland ecotype), Torrey rush (Juncus torreyi), Ehgelmann's spike-rush, caric sedges (Carex spp.), and umbrella sedges (Cyperus spp.). Species of the latter two genera could not be identified in their current vegetative (pre-bloom) phenological stages.

17. Bulrush and cordgrass- Freshwater marsh on margin of salt plains in Central Lowlands physiographic province in northcentral Oklahoma. The dominant range plant was American bulrush (known also as chair-makers' bulrush, chair-makers' rush, and Olney threesquare) with prairie cordgrass (Spartina pectinata) and tapertip flatsedge or tapertip umbrella-sedge (Cyperus acuminatus) as associates. Major forbs were spotted water hemlock (Cicuta maculata) and blue vervain (Verbena hastata). Pink thoroughwort, pink Joe Pye-weed (Eupatorum incarnatum) was a less abundant forb.

This wetland did not typically have an inundated or covered soil surface yearlong, but the soil was saturated most of the year so that this was a marsh, an herbaceous wetland. This freshwater marsh was situated within the overall tallgrass prairie. In the framework of Landscape Ecology, freshwater marshes were patches within the matrix of tallgrass prairie.

Salt Plains National Wildlife Refuge, Grant County, Oklahooma. Late July; peak standing crop (or nearly so). FRES No. 41 (Wet Grasslands Ecosystem). K-42-Tule Mrashes. No SRM designation of a rangeland cover type for tule marshes. In Brown et al. (1998, p. 45) this would be Bulrush Series 242.33 (if one was shown which it was not) of Plains Interior Marshland 242.3. Central Great Plains- Prairie Tableland Ecoregion, 27d (Woods et al., 2005).

18. Botanical players on a wetland stage- Two views of a freshwater marsh in the salt plains area of northcentral Oklahoma showing principal plant species and structure (along with botanical composition). In the first or upper slide American bulrush (background), overall dominant species was shown with tapertip flatsedge (foregound) the associate species.

The second slide featured spotted water hemlock and blue vervain, the two major forbs in the freshwater marsh community.

Salt Plains National Wildlife Refuge, Grant County, Oklahoma. Late July; just about all species at peak standing crop and full bloom. FRES No. 41 (Wet Grasslands Ecosystem). K-42-Tule Mrashes. No SRM designation of a rangeland cover type for tule marshes. In Brown et al. (1998, p. 45) this would be Bulrush Series 242.33 (if one was shown which it was not) of Plains Interior Marshland 242.3. Central Great Plains- Prairie Tableland Ecoregion, 27d (Woods et al., 2005).

19. A major, multi-named, and confusing bulrush- Dense stand (first slide) and flower cluster near end of sexual shoot (second slide) of an aquatic grasslike plant known variously as American bulrush, chairmaker's rush, common threesquare, Olney's bulrush, and Olney's threesquare (among others). These examples were growing in a freshwater marsh in the Salt Plains area of tallgrass prairie in northcentral Oklahoma. This plant was the overall dominant species of this wetland.

The triangular shape of the shoot of this Scirpus species was obvious in the second slide.

Salt Plains National Wildlife Refuge, Grant County, Oklahoma. Late July; fruit-ripe stage of phenology.

20. Taxonomy and nomenclature: Recent (beginning primarily in 1990s) names and interpretations of phylogenetic relations in the bulrushes or tules is a bloody mess. According to the unpublished Great Salt Plains Refuge plant species list (and referencing back to published species synonyms) these species of Scirpus were on this range:. 1) chairmakers' rush, American bulrush, or three square (S. americanus), 2) salt marsh or alkali bulrush (S. paludosus= S. maritimus),and 3) soft-stem or great bulrush (S. validus). S validus was shown on the refuge list as synonyous with S. tabernaemontani which has usually been shown as synonymous with S. acutus which was generally known as a separate species with common names of hard-stem or, also, great bulrush. Thus there might be as many as four species of bulrush or tule on the range of this refuge. Finally, it was noted that all of these tule or bulrush species which were previously shown as Scirpus had been changed to Schoenoplectus.

These examples of chairmakers' rush, American bulrush, or three square were from Cherry County, Nebraska. Late June (near peak biomass, some shoots at full-boom).

21. "Froggie Went A-Courtin'"- Frog (sometimes, fog)-fruit (Phyla lanceolata) is a member of the vervain family (Verbenaceae) that grows on a remarkably diverse array of habitats. These range environments generally seem to be locally disturbed and/or wet microsites. One seemingly inconsistent habitat was as an associate species on a tule marsh in northcentral Oklahoma at the western edge of the the tallgrass prairie region. Specimens of frog-fruit there were not as advanced as others previously photographed by the author so the latter were "transplanted" here. Either way and on both ranges, frog-fruit fared better than Froggie (or was it "Froggy"?) and Miss Mouse in the timeless ballad.

Ottawa County, Oklahoma. June; full-bloom stage.

22. Spotted deadly- Spotted waterlock (Cicuta maculata) shown as whole herbacous view (first slide) and upper shoots complete with inflorescence (second slide) growing in a freshwater marsh in the Salt Plains of northcentral Oklahoma. This plant was growing in association with American bulrush or chairmaker's rush and blue vervain. Larger shoots of all these wetland plants extended to heights of seven (or, rarely, more) feet.

This wetland range community was a botanically diverse (and interesting) assemblage of plants.

Technical note: The second slide was over-exposded due to lens malfunction or, more accurately, due to improper lens maintenance. Unbeknownst to the photographer, an incompetent servicemen had applied white grease as a lens lubricant years earlier when the lens underwent routine preventive maintenance! Yes,"believe it or not" the servicemen were downright stupid and used tool grease in a camera lens. Wonder is that the ole Nikon/Nikoor macrolense had worked for years and thousands of slides before it eventually could no longer perform properly. Fortunately there are competent servicemen and one of these restored the old Nikkor to its original proper functioning. Taken in conjuction with other photographs of spotted water hemlock, students atill get a composite view of spotted watr hemlock.

Salt Plains National Wildlife Refuge, Grant County, Oklahoma. Late July; full-bloom phenological stage.

23. Poisonous on the plains- Upper shoots with umbels (first slide) and compound leaves (second slide) of spotted water hemlock growing in a freshwater marsh in the Salt Plains of northcentral Oklahoma.

Spotted water hemlock is one of the most toxic of North American range plants. In fact, many toxicologists and plant taxonomists regard it as the single most poisonous native to North America. The poisonous principle is cicutoxin, a naturally occuring alcohol. This is a species of large, rank-growing plants that are generally unpalatable to just about everything, However, numerous deaths--to both livestock and humans--have from ingestion of spotted water hemlock. Hence, "cowbane"is one comon name that reflects this toxicity to livestock.

Human deaths are almost always due to accidental poisoning when the immense roots of this member of the parsely family are mistakenly eaten for "carrots" or "parsnips". When handling plants of this species (as in photography of range plants) all body parts that came in contact with them should be washed thoroughly and as soon as possible.

Salt Plains National Wildlife Refuge, Grant County, Oklahoma. Late July; full-bloom phenological stage.

24. Toxic details- Main stem and branches of spotted water hemlock growing in a freshwater marsh in the Salt Plains of northcentral Oklahoma. As is typical of many species of the Umbelliferae, the carrot or parsely family, spotted water hemlock is a biennial or, less commony, a perennial. Also typical of the umbelliferous species are the large, bulbous roots of this virulently poisonous plant.

Salt Plains National Wildlife Refuge, Grant County, Oklahoma. Late July; peak standing crop phenological stage.

25. No spots on flowers- Umbel (upper slide) and clusters of flowers in umbel (seond slide) of spotted water hemlock growing in a freshwater marsh in the Salt Plains of northcentral Oklahoma.

Salt Plains National Wildlife Refuge, Grant County, Oklahoma. Late July; peak standing crop phenological stage.

26. Non-poisonous with poisonous- Upper shoot of blue vervain (Verbena hastata) accompanied by American bulrush and spotted water hemlock growing in a freshwater marsh in the Salt Plains of northcentral Oklahoma.

This is one of the larger, taller Verbena species. It prefers wet to moist habitats like that of this wetland.

Salt Plains National Wildlife Refuge, Grant County, Oklahoma. Late July; full-bloom phenological stage.

27. Blue-topped- Upper shoots (first slide) and flower cluster or inflorescence (second slide) of blue vervain growing in association with American bulrush or chairmaker's rush and spotted water hemlock. Individuals of this species grow to comparatively large size (heights of seven or more feet on this marsh).

Forage value of blue vervain to livestock is essentially zero. It is, however, a good bee plant. In Range Management one never assumes that a species of low palatability to livestock is a weed or competitor to "valuable species". It may be that species like blue vervain provide certain features, factors, or habitat conditions that are beneficial or even essential to more palatable range species.

Salt Plains National Wildlife Reguge, Grant County,Oklahoma. Late July; full-bloom phenological stage.

Marshes of the Nebraska Sandhills

The following section provided a representative sample of wetlands (with herbaceous and/or woody plants) that developed in the semiarid Nebraska Sandhills.

19. 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).

20. 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, p. 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).

21. "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).

22. 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.

23. 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).

24. 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).

25. 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).

26. * "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).

27. 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 arangement.

28. 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.

29. 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.

30. 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. .

31. 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.

32. 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).

Savanna Wetlands (Wetland Savannas, if preferred)

There is a variety of range vegetation that includes both woody and herbaceous, especially grasses and grasslike plants, such that the plant communities cannot be unambiguously described as either swamp or marsh. Rather, such wetland range communities are sevannas or, more completely, wetland savannas. Examples of these savanna wetlands range types include vegetation that is transitional (ecotonal) among bottomland forests (eg. cottonwood [Populus deltoides], hackberry or sugarberry (Celtis spp.]-elm (Ulmus spp.], sycamore [Platanus occidentalis]) willow carrs, etc. and wet prairies, marshes, or a combination of these herbaceous wetlands. These ecotonal wetland range types are present in precipitation zones ranging from wet to arid. Examples follow immediately below.

33. Savanna wetland in the subhumid zone- Synopsis views of a combination eastern cottonwood-peachleaf willow wet woodland and a tule (bulrush)-cattail-umbrella or flat sedge marsh in the Smokey Hills which are part of the border of High Plains and Central Lowlands physiographic provinces known as the Plains Border physiographic section (Frye and Swineford, 1949). This was the western variety of eastern cottonwood (Populus deltoides var. occidentalis) as mapped by McGregor et al. (1977), a conspicuous individual of which was the tallest woody plant in both of these slides. Weber (1990) described the western taxon of plains or eastern cottonwood as Populus deltoides subsp. monilifera).

The tule was soft-stem or great bulrush, (Scirpus validus), the flat sedge was tapertip flat (umbrella) sedge (Cyperus accumulatus), and the sole member of Typhaceae was narrowleaf cattail (Typha domingensis).

Range vegetation in the immediate foreground was not part of the wetland (or at least only on the margins of it), but was typical tallgrass prairie comprised mostly of a bottomland ecotype of switchgrass (Panicum virgatum) and tall dropseed (Sporobolus asper subsp. asper). The prominent forb on this margin was Illinois bundleflower (Desmanthus illionensis).

Ellsworth County, Kansas. Mid-July, early estival aspect. Salicaceae form of savanna with marsh with closest range ecosystem and type being FRES No. 417 (Wet Grasslands Ecosystem). K-42 (Tule Marshes). No SRM for the wetland (601 or 710 for tallgrass paririe). Woodland element was variant of SAF 63 (cottonwood). Combination of Plains and Great Basin Riparian Scrub biotic community, 222.2, Cottonwood-Willow Series, 222.21 and Plains Interior Marshland biotic community 242.3, Sedge Series, 242.31 of (Brown et al., 1998, p. 44). Central Great Plains- Smoky Hills Ecoregion, 27a (Chapman et al., 2001).

34. Into the "swamp" in the Smoky Hills- Series of sequential photo-points of a savanna wetland moving consistetly closer to the woody element of eastern cottonwood and peachleaf willow. Although there was some naturalized smooth brome (Bromus inermis) close to the shaded habitat of trees, the herbaceous component of this patch of savanna in a matrix of tallgrass prairie (description in terms of Landscape Ecology) was a locally varying community of great or soft-stem bulrush, tapertip flat sedge, and narrowleaf cattail. These three native species of grasslike plants intermingled in differing proportions as well as growing in local colonies or populations of their singular species.

The conspicuous forb with its dormant shoot of ripe, shedding fruits was pale or dock (Rumex altissimus). The other forb (not visible in these photographs) was shown below.

A characteristic trunk of peachleaf willow was presented in the third photograph. This and black willow are the only large willows in Kansas (Stephens , 1969, ps. 22-23, 26-27). Both Salix species typically have leaning trunks with very similar bark, but peachleaf willow can be distinguished by the pale or whitish undersides of its leaves (Stephens, 1969, ps. 23, 27).

Ellsworth County, Kansas. Mid-July, early estival aspect. Salicaceae form of savanna with marsh with closest range ecosystem and type being FRES No. 41 (Wet Grasslands Ecosystem). K-42 (Tule Marshes). No SRM for the wetland (601 or 710 for tallgrass paririe). No FRES unit seemed appropriate for the woody component of this vegetation. Woodland element was variant of SAF 63 (Cottonwood). Combination of Plains and Great Basin Riparian Deciduous Forest biotic community, 222.2, Cottonwood-Willow Series, 222.21 and Plains Interior Marshland biotic community 242.3, Sedge Series, 242.31 of Brown et al., (1998, ps. 43, 45). Central Great Plains- Smoky Hills Ecoregion, 27a (Chapman et al., 2001).

35. Grasslike plants on a savanna wetland- Herbaceous component of a cottonwood-willow-marsh savanna in the Smoky Hills tallgrass-true prairie region dominated locally by tapertip flat (umbrella) sedge with great or soft-stem bulrush and narrowleaf cattail as associate species. The Cyperaceae and Typhaceae were both represented while Poaceae (Gramineae) was excluded from this community and relegated to the margins of the marsh (shown and explained in the immediately preceding slides and captions).

Details of a local stand of tapertip flat sedge with scattered shoots of soft-stem bulrush were presented in the second photograph. On other local spots (microsites or microhabitats) of the marsh vegetation soft-stem or great bulrush was dominant (see very next slide below).

This wetland range ecosystem was in the Plains Border physiographic section (Frye and Swineford, 1949), a sort of "no man's land" between the High Plains and Central Lowlands physiographic provinces, of the continental interior of North America.

36. Our turn- Sward of the marsh component of a savanna wetland in the Smoky Hills within the tallgrass-true prairie types region of the Plains Border physiographic section (Frye and Swineford, 1949). Great tule or soft-stem bulrush was the dominant with annual marsh elder or sumpweed (Iva annua), the only forb of consequence and consistency on most of this marsh, the associate species. Tapertip flat sedge, overall dominant of the marsh portion, was present as a lower-herbaceous level species in this local assemblage.

Ellsworth County, Kansas. Mid-July, early estival aspect. Herbaceous element of a savanna wetland that was described by FRES No. 41 (Wet Grasslands Ecosystem). K-42 (Tule Marshes). No SRM. Plains Interior Marshland biotic community 242.3, Sedge Series, 242.31 of Brown et al. (1998, p. 45). Central Great Plains- Smoky Hills Ecoregion, 27a (Chapman et al., 2001).

37. My turn- "Center stage" in this slide went to narrowleaf cattail which on this savanna wetland in the Kansas Smoky Hills was typically an associate to the overall dominant, tapertip flat sedge. On this particular wetland range narrowleaf cattail grew as a population (single species stand) only rarely. More commonly either narrowleaf cattail or great bulrush was the associate species to the dominant tapertip flat sedge.

38. Bulrushes by willows- Local stand of great or soft-stem bulrush (Scirpus validus) by peachleaf willow on a savanna wetland in the Smoky Hills. This bulrush colony largely excluded the other major species (narrowleaf cattail, tapertip flat sedge, sumpweed) of this wetland range. These shoots were at the ripening fruit stage.