Miscellaneous Grasslands

Glades, Balds, and Barrens

Chert Glade

1. Ozark Chert Glade- Glades makeup one of several kinds of range "barrens". Nestled among the oak-hickory forest and the tallgrass prairies in the Ozark Plateau are small areas of rock outcrops or shallow, rocky soils on hilltops, and other similar kinds of land and its vegetation known as "glades". In other words, within this southern portion of the Prairie Peninsula there are locally restricted sites that are edaphic or topographic climaxes within the climatic climax of the western extension of the Eastern Deciduous Forest and the Tallgrass Prairie Region. Nelson (1985, p. 99) defined glades as "rocky barrens dominated by a herbaceous flora with sparse woody vegetation". This fits the standard definition of savanna. In fact, savannas and barrens traditionally have been considered as similar environments (see especiallly Anderson et al., 1999). Nelson (1985, p. 99) further specified that glades "are topographically located on moderate to steep slopes in deeply dissected drainages or hilly to mountainous terrain, and usually have a southern or westen exposure". Bedrock is at or near the soil surface and the shallow soils are subject to frost heaving in winter and are very dry during much of the frost-free growing season following soil saturatrion during spring, sometimes fall.

Glades or, as they are more commonly known by the local hillbillies, "balds" or "bald knobs" are one form of "barrens". Glades are classified as to their parent material. Missouri has greater diversity in balds than any state and has limestone, dolomite, chert, sandstone, shale and igneous glades. The bald seen here in the Ozark Highlands of southwestern Missouri is a chert glade derived from massive brecciated chert of the Elsey Formation (Nelson, 1985, p. 106). An ephemeral (ie. "wet-weather") stream flows through the center draw in early to mid-spring and there are seeps from under the chert outcrops which support the"runt" or scrub form of blackjack and post oaks (Quercus marilandica and Q. stellata) and common "hackberry (Celtis occidentalis). The domiant grass with the "lacy" or "dainty" appearance in the center and left-center foreground is Festuca paradoxica. The dominant yellow composite is tickseed coreopsis (Coreopsis lanceloata). This is the bottom of the bald which supports the vernal, cool-season, most mesophytic species. The surrounding oak-hickory forest was described immediately below. Wildcat Glade Natural Area, Newton County, Missouri. Vernal aspect, May. FRES of the glade vegetation is 39 (Prairie Ecosystem), K-66 (Bluestem Prairie), FRES of the neighboring forest is No. 91 (Oak-Hickory Forest), and FRES of the combined glade and forest vegetation is No. 73 (Mosaic of Numbers 66 and 91). SRM 803 (Missouri Glades). Range site is Limestone Rock Outcrop. Northwestern Great Plains-Semiarid Pierre Shale Plains Ecoregion, 43g ( Bryce et al., undated).

2. Mid-slope vegetation on an Ozark Highlands chert glade (= chert barrens)- Above the mesic habitat of the draw in the chert glade seen in the previous slide is mid-slope vegetation dominated by big bluestem (Andropogon gerardii) and an uplant ecotype of switchgrass (Panicum virgatum). The scrub trees are blackjack oak, post oak, and hackberry. These same oak species grow in the surrounding oak-hickory forest which, growing interspersed with tallgrass prairie (the Prairie Peninsula), is the climatic climax vegetation. The forest has as its dominant, however, black oak (Quercus velutina) which besides post and blackjack oak has as associates black hickory (Carya texana), red or sweet pignut hickory (C. ovalis), and mockernut or white hickory (C. tomentosa). There were a few American elm (Ulmus americana) and even hackberry (Celtis occidentalis).Redbud (Cercis canadensis) and flowering dogwood (Cornus florida) are dominant shrubs of the adjacent forest. Smooth sumac (Rhus glabra) and small plants of persimmon (Diospyros virginiana) were also present at forest edges and in the glade proper. Glade vegetation has variously four to five layers: 1) tallgrass species like big bluestem and switchgrass, 2) scrub blackjack and post oak trees of single stems but no taller (usually shorter) than the mature tallgrasses such as to be a shrub layer, 3) a mid-height forb and grass layer (eg. composites, festucoid grasses), 4) xerophytic low herb-shrub layer (short forbs, ferns, cactus; especially common on chert outcrop ledges), and 5) xerophytic soil and rock surface layer (cryptogams). Layer 5 is not present at all microsites.

Wildcat Glade Natural Area, Newton County, Missouri. Estival aspect, August. FRES of the glade vegetation is No. 39 (Prairie Ecosystem),K-66 (Bluestem Prairie) FRES of the oak-hickory forest is No. 91, and FRES of the combined glade and forest vegetation is No.73 (Mosaic of Numbers 66 and 91). SRM 803 (Missouri Glades). Range site is Limestone Rock Outcrop. Northwestern Great Plains-Semiarid Pierre Shale Plains Ecoregion, 43g ( Bryce et al., undated).

Chert Glade societies- In context of Plant Ecology, specifically Plant Community Ecology or Synecology, the term society refers to "a community characterized by one or more subdominants, i.e. species of different life form from those of the regional dominants" (Weaver and Clements, 1938, p. 96). In this regard there are two kinds of vegetational societies: 1) seasonal aspects and 2) layering. Both of these kinds of society are important and distinctive in chert glades, but it is s the seasonal or aspect society that was addressed and used as a teaching construct in this treatment of chert glades as range vegetation.

Glades (chert, dolomite, granite, etc.) probably have greater differences in seasonal composition (more variation in the annual cycle of range plant life) as seen from one vegetational society (seasonal society) to another than any natural plant community in North America. This range vegetation varies from a primarily thallophytic (lichens)-bryophytic (mosses) winter (hibernal) society to a forb-dominated spring (vernal) society to a grass-dominated summer (estival)/autmmn (autumnal) society. Given that a natural plant community is classified, grouped, titled, etc. on a yearlong basis and based on the dominant plant growth form, family, or category it would be most correct (or most nearly correct) to regard glades as grasslands.

Grasses (major or dominant tallgrass prairie grass species at that) are the dominant plant family or range plant group of glade vegetation at peak standing crop. In fact, plant species composition of chert and dolomite glades or bald knobs in the Ozark (Springfield) Plateau is about the same as tallgrass prairie in the Flint Hills and Osage Cuestas. The difference in range vegetation of chert galdes being described here and rock-underlaid tallgrass prairie as in the Flint Hills is that some vegetational societies in the glades are dominated by forbs or nonvascular plants to a much greater degree than are seasonal societies of tallgrass prairie vegetation in the Flint Hills or non-glade prairies or prairie meadows in the Ozark Plateau are dominated by grasses (cool-season or warm-season; winter dormancy or autumnal peak biomass).

The discussion of range vegetation in the chert glades of the Ozark Plateau was organized as to the seasonal societies of this natural plant community.

Chert Glade in Spring

Range vegetation of Ozark Plateau chert glades in the spring--the vernal society of this natural plant community--is dominated by forbs. In fact, the spring vegetation of chert glades could be descriptively categorized as forbland Of course, on a annual or dominant (as to cover and herbage yield) plant basis, glades are grasslands, specifically tallgrass prairie. Presented below were images and descriptions of range plant communities that developed on chert glades in the Ozark (Springfield) Plateau.

The first part of Chert Glade in the Spring showed the overall grassland or grassland-hardwood savanna with emphasis on physiogonomy and structure. These views of the general range plant community were followed by coverage of the vernal society (ies) on the shallower, rockier haabitats. Range vegetation on these more harsh (even by chert glade standards) environments is more colorful and plant species-diverse--as it is dominated by forbs--than on the "more hospitable" edaphic habitats that afford better developed, deeper soils with greater water-holding capacity and thereby are dominated by perennial prairie grasses.

Big picture of a little area- Chert dominated by big bluestem and switchgrass overall, but with starved, dwarf, or poverty panicgrass (Panicum depauperatum) as a local dominant with local abundance of rough or secret dropseed (Sporobolus s clandestinus) plus western yarrow (Achillea lanulosa), yellow stonecrop (Sedum nuttallii), three-flower melic (Melica nitens), goat's rue Tephrosia virginicca), tickseed or lanceleaf coreopsis (Coreopsis lanceolata) roundhead lespedeza (Lespedeza capitata), clasping Venus's looking glass (Specularia perfoliata), Hairy lip-fern (Cheilanthes lanosa), little spike moss (Selaginella rupestris), and numerous thallophytes among the herbaceous range plants.. (Hardwood trees and shrubs have their entrances below.)

Glade Natural Area, Newton County, Missouri. Vernal aspect, Early June. FRES of the glade vegetation is No. 39 (Prairie Ecosystem),K-66 (Bluestem Prairie) FRES of the oak-hickory forest is No. 91, and FRES of the combined glade and forest vegetation is No.73 (Mosaic of Numbers 66 and 91). SRM 803 (Missouri Glades). Range site is Limestone Rock Outcrop. Northwestern Great Plains-Semiarid Pierre Shale Plains Ecoregion, 43g ( Bryce et al., undated).

At the edge of woods- View of a tallgrass (big bluestem and switchgrass, mostly)-dominated chert glade surrounded by oak-hickory forest (at least before the whiteman took down most of the forest for housing and recreational development). Woody species have invaded local microsites of the grassland as shown below. It cannot be ascertained with certainly whether such woody invasion by native hardwoods is a historic natural event with invasion limited by drought, natural (lightening-ignited) or Indian-set fires or whether this was accelerated by adjacent human activities. At any rate, chert glades are a unique--very spatially restricted--form of tallgrass prairie or, perhaps, tallgrass-hardwood savanna.

Herbaceous species shown here are the same as those listed immediately above and below. Adjacent major hardwood trees were blackjack oak, post oak, and black oak with numerous hickory species that were listed in the introductory slide captions.There were also some plants of American elm, persimmon, hackberry, redbud, and smooth sumac.

Natives, but were they visitors or permanent residents?- Two views of woody plant invasion of big bluestem-switchgrass dominated tallgrass prairie on chert glades in the western Springfield Plateau section of the Ozark Plateau (Highlands) in southwestern Missouri. The first or top slide featured an American elm (llargest woody plant in center) that was showing burnt back leaves from heat (drought and temperture) stress. The small elm sapling was accompanied with blackjack oak, persimmon, smooth sumac, and a small hackberry. The second slide featured a persimmon sapling (right foreground) and with blackjack oak, American elm saplings, and small patches of smooth suma. In addition to the two tallgrass dominants, three-flower melic, rough dropseed, dwarf or secret panicgrass, lanceleaf tickseed or coreopsis, rounghead lespedeza, western yarrow, clasping Venus lookingglass, little spike moss, and hairy lip-fern were other herbaceous species.

At time of photograph, all these woody plants had survived two growing seasons (not consecutive years) in which Extreme Drought existed for protracted periods Plus, other recent growing seasons had been abnormall dry. In other words, so far bad droughts had not killed off the woody species. "Time will tell", including whether drought or drought plus fire is necessary to maintain the tallgrass prairie on the chert glades.

3. Chert glade sampler- Climax range plant community on an outcrop of chert in the western Ozark (Springfield) Plateau. The vernal society of this unique range vegetation included Ohio spiderwort or bluejacket spiderwort (Tradescantha ohioensis); yellow or Nuttall's stonecrop or golden sedum (Sedum nuttallianum), lanceleaf tickseed (Coreopsis lanceolata), sheep's sorrel or red sorrel (Rumex acetosella); hairy lipfern (Cheilanthes lanosa); puffsheath dropseed (Sporobolus neglectus); poverty dropseed (S. vaginiflorus); winter bentgrass (Agrostis hyemalis); a cool-season annual associate; and, at early spring growth stage, switchgrass (Panicum virgatum) which is often the climax dominant of this range community at peak standing crop stage. The major nonvascular plant in this "phototransect" was a foliose lichen, Cumberland xanthoparmelia lichen (Xanthoparmelia cumberlandia).

Slopes just above Shoal Creek, a stream the size of a small river, in the Springfield Plateau.

Wildcat Glade Natural Area, Newton County, Missouri. Late May; vernal aspect. FRES No. 39 (Prairie Ecosystem), K-66 (Bluestem Prairie). SRM 803 (Missouri Glades). Closest biotic community unit in Brown et al. (1998, p. ) was Cold Temperate Grassland 142, Plains Grassland 142.1, Bluestem "Tall-Grass" Series 142.11. Range site is Limestone Rock Outcrop (Soil Conservation Service, 1989 did not list range sites for Newton County, Missouri) or Very Shallow with limestone flagstones for adjoining Ottawa County, Oklahoma (Soil Conservation Service, 1964). Ozark Highlands- Springfield Plateau ecoregion 39a (Chapman et al., 2002).

4. Glory of the glades- The vernal society of native vegetation of a chert glade dominated by forbs such as Ohio spiderwort, yellow or Nuttall's stonecrop, lanceleaf tickseed, hairy lip-fern, and red or sheep sorrel. Grass species included three-flower melic (Melica nitens), the dominant grass at this season (in this seasonal society); winter bentgrass, a cool-season annual associate; switchgrass; puffsheath dropseed; and poverty dropseed.

Woody species included blackjack oak (Quercus marilandica) as shrub-sized trees and eastern pricklypear (Opuntia compressa= O. humifusa). Dominant lichen on chert outcropping in these two photographs was primarily the foliose lichen, Cumberland xanthoparmelia lichen (Xanthoparmelia cumberlandia). Grass species in the vegetation presented in these two photographs included three-flower melic (Melica nitens), the dominant grass at this season (in this seasonal society); winter bentgrass, a cool-season annual associate; switchgrass; puffsheath dropseed; and poverty dropseed.

The large Ozark Plateau stream, Shoal Creek, was in the background of these beautiful spring scenes of the chert glades.

5. Diversity and splendor amid the rocks- Spring vegetation on a chert glade range in the Ozark Plateau. Forbs were dominant in this vernal society with grasses the second major group of range plants. Forb species included Ohio spiderwort, yellow or Nuttall's stonecrop, lanceleaf tickseed, hairy lip-fern, and red or sheep sorrel. Grasses included three-flower melic, the dominant grass at this season (in this seasonal society); winter bentgrass, a cool-season annual associate; switchgrass; puffsheath dropseed; and poverty dropseed.

There were a few plants of eastern pricklypear. the main thallophyte growing on the chert outcroppings was a foliose lichen, Cumberland xanthoparmelia lichen.

Upper slope just above Shoal Creek in southwestern Missouri.

6. Forage and beauty on the outcrops- Vernal society of climax vegetation of an Ozark Plateau chert glade. Major range plant species were forbs including Ohio spiderwort or bluejacket, yellow or Nuttall's stonecrop, lanceleaf tickseed, hairy lip-fern, and red or sheep sorrel.

The main range plant growing on the chert outcroppings was a thallophyte, a foliose lichen, Cumberland xanthoparmelia lichen. The major grasses on this chert glade were switchgrass, the overall dominant species of this range plant community, which was visible in these two "photplots". Other grasses on this glade which were not visible included starved panicgrass, poverty dropseed, puffsheath dropseed, and winter bentgrass.

The only woody plant species present was eastern pricklypear.

These rock outcrops were on the upper slope above Shoal Creek, a stream the size of a small river, in the western Ozark Plateau in southwestern Missouri.

7. Botanical details on the chert- Range vegetation on chert outcrops (flagstones) in an Ozark (Springfield) Plateau. Most of the plant species growing in the interspaces among these rocks were forbs including lanceleaf tickseed, hairy lip-fern, Ohio spiderwort or bluejacket, yellow or Nuttall's stonecrop, and red or sheep sorrel. The first slide was a composite view of this botanical assemblage.

The second slide included the two vascular plant species of switchgrass, the dominant range plant of the chert glades, (the long, linear-leafed clumps of herbage) and hairy lip-fern (the shorter, broad-leafed clumps). Chert outcrops were covered by vrious lichen species the dominant of which was the foliose lichen, Cumberland xanthoparmelia lichen.

8. A cherty home- Three "photoquadrants" of spring vegetation growing on (mostly spaces amid) chert flagstones on a glade in the western Ozark Plateau. Forbs were the major group of native range plants in these three shots. The first view was at the edge of a chert outcrop with Cumberland xanthoparmelia lichen, a foliose lichen, growing on the edge of a chert flagstone and hairy lip-fern growing in shallow soil at the edge of this outcrop. The background in this first slide was vegetation comprised primarily of switchgrass with some cover of the rosette panicgrass known as starved, dwarf, or poverty panicgrass (Panicum depauperatum) .

The second slide was also of hairy lip-fern with switchgrass growing out of clumps of the fern and with some plants of Nuttall's or yellow (golden) stonecrop and lanceleaf tickseed or lanceleaf coreopsis in the right upper corner while Cumberland xanthoparmelia lichen "stood sentry" on chert flagstones. The third slide was a closer-in view of plant species presented in the second slide (ie. lanceleaf tickseed, switchgrass, hairy liip-fern, yellow or golden stonecrop and Cumberland xanthoparmelia lichen "on the rocks".

These flagstones werre chert outcrops on the slope above Shoal Creek, a larger creek in the western Springfield Plateau in southwestern Missouri.

Wildcat Glade Natural Area, Newton County, Missouri.Late May; vernal aspect. FRES No. 39 (Prairie Ecosystem), K-66 (Bluestem Prairie). SRM 803 (Missouri Glades). Closest biotic community unit in Brown et al. (1998, p. ) was Cold Temperate Grassland 142, Plains Grassland 142.1, Bluestem "Tall-Grass" Series 142.11. Range site is Limestone Rock Outcrop (Soil Conservation Service, 1989 did not list range sites for Newton County, Missouri) or Very Shallow with limestone flagstones for adjoining Ottawa County, Oklahoma (Soil Conservation Service, 1964). Ozark Highlands- Springfield Plateau ecoregion 39a (Chapman et al., 2002).

9. Ones who thrive on the outcrops- Three major botanical players in the vernal society of natural vegetation growing in spaces of shallow soil among chert flagstones in a Ozark Plateau glade. The long narrow leaves were those of switchgrass, the climax dominant of this general range plant community. In these slides the greatest cover in this vernal aspect (society) was hairy lip-fern charactterized by its comparatively greater size and density of shoots with the compound fronds. The low yellow plants (eg. left foreground of first slide( was yellow or golden stonecrop or, also Nuttall's stonecrop (sedum). Yellow stonecrop has the Crassulacean Acid Metabolism pathway, the most water-efficient of the three photosynthetic pathways. Switchgrass has the C4 pathway which uses water more efficiently than the C3 pathway characteristic of cool-season species.

Cumberland xanthoparmelia lichen, a foliose lichen, was the major thallophyte on chert flagstones in these two views.

10. Home is where the chert is- Some dominants of the vernal society of the climax range vegetation on a chert glade in the western Ozark Plateau. The "featured speaker" was lanceleaf tickseed or lanceleaf coreopsis with entire shoots shown in the first slide while basal shoot portions were emphasized in the second slide along with shoots of yellow or golden stonecrop (or Nuttall's sedum), hairy lip-fern, and some species of moss. There were also plants of bluejacket or Ohio spiderwort, hairy lip-fern, Nuttall's stonecrop, starved or poverty rosette panicgrass, and switchgrass in the first slide. Cumberland xanthoparmelia lichen, a foliose lichen was growing on chert outcrops in both slides.

Bracted on chert- Bracted platain (Plantago aristata) seen as the entire aboveground portion (first slide) and three flower clusters (second slide). This Plantago species is a native and not one of those whiteman-introduced Eurasian weeds which are not worthy to be mentioned otherwise in the company of this native that was clinging to life in a rare chert glade at the southwestern prairie Peninsula (Transeau, 1935) in the Springfield Plateau of southwest Missouri. Ironically, this annual with a species range extending to New England, has itself become an introduced weed in Europe and the British Isles.

An excellent online source for extraordinary closeup photographs of this and other plants of the Show Me State is: Missouriplants.com.

Wildcat Glade Natural Area, Newton County, Missouri. Early June, peak flowering phenological stage.

11. Depauperate but not impoverished- Starved, dwarf, or poverty panicgrass (Panicum depauperatum) was a locally dominant species in the vernal society of the range vegetation of a chert glade in the Ozark (Springfield) Plateau. This is one of the rosette panicgrass species that Hitchcock and Chase (1950) placed in the section, Dichanthelium, hence Dichanthelium depauperatum according to some authors. These rosette panicgrasses are cool-season species with the C3 photosynthetic pathway. Starved rosette pancigrass is a cespitose (bunchgrass) species.

Note that P. depauperatum is a local dominant of one of the four seasonal societies or aspects and not an overall or yearlong, community wide dominant.

Switchgrass (Panicum virgatum) is an overall dominant species of chert glades, but this dominance is in the estival and autumnal societies and at peak standing cropSwitchgrass is one of the four major tallgrass species of tallgrass prairie: the Four Horsemen of the Prairies. Switchgrass is a large, robust warm-season species with the C4 photosynthetic pathway.

Location note: slides of plant species found in the above and below images of vegetation were presented at the end of the end of the subsection, Chert Glade in Summer.

The second part of Chert Glade in the Spring featured the vernal society (ies) on the larger areas that have soils with greater development of their profiles and, hence, are deeper so as to offer a rhizosphere,"the zone of soil immediately adjacent to plant roots in which the kinds, numbers, or activities of microorganisms differ from that of the bulk soil" (Soil Science Society of America, 2001), soil water features, and, most likely, soil fertility that are nmore favorable for those plant species which have--and depend on--larger root systems (especially, native perennial grass species).

Chert Glade in Summer

This sub-section presented images and descriptions of the summer society of range vegetation on a chert glade in the Ozark Plateau. The vernal and autumnal aspects and their vegetational societies merge slowly into those of the vernal or summer portion of the annual cycle of life on the edaphically harsh habitat of the chert glades.

12. Early summer and already drying out- Sample of a local community of range plants on the shallow, rocky soil of a chert glade in the western Ozark Plateau shortly after onset of the summer portion of annual life cycle of this range vegetation. The local dominant in this "photoplot" was a holdover forb of the vernal society that was loclly dominant. Whorled milkweed (Asclepias verticillata) was the dominant of this microsite at this point in the annual cycle of plant life. The associate species at this time was starved, poverty, or dwarf rosette panicgrass.

The ultimate end-of-growing season (stage peak herbage yield= peak standing crop) dominant in the overall range plant community of the chert glades was switchgrass. For now, whorled milkweed was holding forth as prince of this rock prairie.

Wildcat Glade Natural Area, Newton County, Missouri. Early July. FRES No. 39 (Prairie Ecosystem), K-66 (Bluestem Prairie). SRM 803 (Missouri Glades). Closest biotic community unit in Brown et al. (1998, p. ) was Cold Temperate Grassland 142, Plains Grassland 142.1, Bluestem "Tall-Grass" Series 142.11. Range site is Limestone Rock Outcrop (Soil Conservation Service, 1989 did not list range sites for Newton County, Missouri) or Very Shallow with limestone flagstones for adjoining Ottawa County, Oklahoma (Soil Conservation Service, 1964). Ozark Highlands- Springfield Plateau ecoregion 39a (Chapman et al., 2002).

13. Detail view of xerophytic layer of low herbs-shrubs on a rock outcrop ledge on a chert bald or barrens in the Ozark Plateau- Hairy lip fern (Cheilanthes lanosa) is the dominant species of most of the microsites of soils occurring between ledges of chert outcropping. Prickly pear (Opuntia compressa) and Missouri black-eyed susan (Rudbeckia missouriensis) are associates. Note extreme desication of the fern nearing end of summer. This is an exception to the usual condition where most fern species are associated with mesic, shaded habitats (in the Ozark Mountains such environments are most common on north and east slopes of bluffs above creeks and rivers). Here a species of fern is a xerophyte and a neighbor to a species of cactus.

Estival aspect, August. Wildcat Glade Natural Area, Newton County, Missouri. FRES No. 39 (Prairie Ecosystem), K-66 (Bluestem Prairie). SRM 803 (Missouri Glades). Range site is Limestone Rock Outcrop.

14. Ground surface of a chert barrens (= chert glade)- Except for the five depauperate big bluestem plants, the plant life on the soil surface of this chert glade in the Ozark Mountains are nonvascular cryptogams. Cryptogams are plants that reproduce sexually by spores (ie. they are not seed plants or spermatophytes). Vascular (having vessels which convey fluids, specifically having the vascular tissues of xylem and phloem that function as conduits through roots, stems, and leaves) plants that reproduce by spores such as ferns, horsetails or scouring rushes, club mosses, and spike mosses are known as pteridophytes (Pteridophyta is the Division of the Plant Kingdom made up of pteridophytes). Pteridophytes are often known generically as "the ferns and fern allies". Tracheophyta is in some plant classification/taxonomic schemes the Division of the Plant Kingdom encompassing both the former Divisions of Pteridophyta and Spermatophyta. In some other taxonomic arrangements the Tracheophyta is elevated to the Kingdom designated Metaphyta, or sometimes, the Plantae. In the taxonomic schemes that use Tracheophyta (the tracheophytes), the Bryophyta (bryophytes are the mosses) and Thallophyta (thallophytes include the algae and fungi) along with the Pteridophyta are the cryptogams with the pteridophytes being vascular cryptogams and the thallphytes and bryophytes being nonvascular cryptogams. The spermatophytes (the gymnosperms and the angiosperms) are designated phanerogams. Embryophtes are those plants in the taxon designated Embryophyta (plants distinguished by having a zygote that develops into a multicellular embryo while in the female sex organ or in the embryo sac) which includes the Bryophyta and the Tracheophyta.

Many of these terms are of the older, classical, or traditional systems that viewed all organisms as either plant or animal. This view was an oversimplification and less precise than the contemporary five kingdom system, but like most of that "generation" of knowledge it was practical and utilitarian for professions and disciplines in the Agricultural Sciences.

The plant community ("plant" by the classical, traditional view) or layer of plant community seen here and in the preceding layer dominated by a species of fern includes spermatophytes, pteridophytes, and thallophytes. Dominant plants of this soil-rock layer are lichens (foliose, fructicose and crustose species); a species of little spike moss is the associate. Observe that the dominant spermatophyte of the prairie, big bluestem, "held its own" even though dwarfed. On this xeric microsite this is the entire plant community: the scrub tree layer is absent as is most of the tallgrass layer. This xeric community (or soil-rock layer) has already progressed phenologically to the autumnal aspect even though it is only August and the rest of the chert glade vegetation is still at the estival aspect. Wildcat Glade Natural Area, Newton County, Missouri. SRM 803 (Missouri Glades). Range site is Limestone Rock Outcrop.

15. Composite shot of the soil-rock community (or, perhaps more specifically, the nonvascular cryptogam vegetation layer) of a chert barrens in the Ozark Plateau- This is a textbook example of a cryptogamic soil surface. The dominant plants are lichens, one of those miraculous kinds of mutualism which like ruminants and their rumen microbes, legumes and their nitrogen-fixing bacteria, and pollinating plants and their pollinators are essential for functioning of range ecosystems and the profession of Range Management.

Lichens were defined by Allaby (1998) as: "a type of composite organism, which consist of a fungus (the mycobiont) and an alga or cyanobacterium (the phycobiont) living in symbiotic [mutualistic] association. A lichen thallus may be crust-like (crustose), scaly or leafy (foliose), or shrubby (fruticose), according to the species."

--Thallus- "a primitive type of vegetative plant body that is not differentiated into stems, leaves, and roots, although analogous structures may be present [used mostly for nonvascular cryptogams]" (Allaby, 1998).

Lichens are major range forage plants on certain types of range in the Arctic Region where they are the mainstay for the caribou/reindeer. On this chert glade they are interesting plants contributing more to soil formationH than to animal diets. Lichens visible include Acarospora chlorophana, Cladonia caroliniana, and Xanthoparmelia spp.). Little spike moss is growing (was growing and is now desicated and dormant) in the cracks among the lichens. Autumnal aspect, August (on this xeric microsite plant phenology has passed to dormancy for the little spike moss and near-dormancy for the lichens). Wildcat Glade Natural Area, Newton County, Missouri. SRM 803 (Missouri Glades). Range site is Limestone Rock Outcrop (this is a microsite thereof).

16. Cladonia caroliniana- This is a fruticose lichen. Fruticose is a lichen growth form that is shrubby or hair-like; the thallus (lichen body) is made up of round or flattened branches. Lichens are some of the first pioneer plants to start the process of plant succession on the parent material of a sere, a xerosere in this instance. This fruticose species is growing on a xeric microsite of a Limestone Rock Outcrope range site. SRM 803 (Missouri Glades). Wildcat Glade Natural Area, Newton County, Missouri. August.

17. Little spike moss (Selaginella rupestris) in the soil-rock layer of a chert barrensin the Ozark Highlands of southwestern Missouri- Numerous spore-bearing cones or strobilii are visible. This colony is growing on a less xeric microsite than that seen in the soil-rock community above and the plants are completing their annual growth cycle at end of summer. Estival aspect, August. Wildcat Glade Natural Area, Newton County, Missouri. SRM 803 (Missouri Glades). Microsite of a Limestone Rock Outcrop range site.

18. Summer synopsis of a chert outcrop- On a flagstone of chert in an Ozark (Springfield) Plateau chert glaade a Cumberland xanthoparmelia lichen (Xanthoparmelia cumberlandia) and dry rock moss or silver beard moss (Grimmia laevigata) had staked out their homesites while hairy lip-fern grew in the shallow soil just below the water-sheeding chert outcrop. Three species so three ecological niches, but the difference between the niche of the lichen and the niche of the rock moss was not apparent to the rock-stepping visitor. Still, the educated rangeman knows the rule: "only one species per niche: the ecological niche defines the species and vice versa".

These three species represent part of the botanical diversity of plant life on chert glades range. The lichen is a thallophyte, a member of the Thallobionta or Thallophyta (subkingdom of the traditional Plantae or classic plant kingdom whose members have no clearly distinctionable roots, stems, or leaves and they produce no seeds). The moss and lip-fern are each an embryophyte, members of the Embryobionta or Embryophyta (subkingdom of the traditional Plantae whose members produce enclosed embryoes within archegonium, spores, or seeds). The rock moss is further classified taxonomically as being a bryophyte (from the Plantae division of Bryophyta). The lip-fern is a pteridophyte (of the Plantae division, Pteridophyta whose members have vascular tissue along with distinguishable roots, shoots (stems and leaves) and reproduce sexually by means of spores). Although the fern produces spores (is a pteridophyte) it is also a vascular plant (possesses the specialized conducting tissues of xylem and phloem) and, therefore, is a tracheophyte (a member of the Plantae subkingdom of Tracheophyta).

There you go, one lessons in Range Ecology and one lesson in Range Plants and both from God's own rock garden.

Wildcat Glade Natural Area, Newton County, Missouri. Late July. FRES No. 39 (Prairie Ecosystem), K-66 (Bluestem Prairie). SRM 803 (Missouri Glades). Closest biotic community unit in Brown et al. (1998, p. ) was Cold Temperate Grassland 142, Plains Grassland 142.1, Bluestem "Tall-Grass" Series 142.11. Range site is Limestone Rock Outcrop (Soil Conservation Service, 1989 did not list range sites for Newton County, Missouri) or Very Shallow with limestone flagstones for adjoining Ottawa County, Oklahoma (Soil Conservation Service, 1964). Ozark Highlands- Springfield Plateau ecoregion 39a (Chapman et al., 2002).

Summer thallophytes and bryophytes- Cover of green byrum moss (Byrum ) and thorn lichen or thorn pixie lichen (Cladonia )

Wildcat Glade Natural Area, Newton County, Missouri. Early June.

Thorny cryptogam- Details of thorn cladonia lichen growing on a chert boulder on a chert glade in the western Ozark (Springfield) Plateau in southwest Missouri.

Wildcat Glade Natural Area, Newton County, Missouri. Early June.

Beards on the rocks Dry rock moss or silver beard moss (Grimmia laevigata)

Wildcat Glade Natural Area, Newton County, Missouri. Early June.

19. Just below the big rocks- On lower slopes of the chert glades (where they "peter out" into an oak-hickory-tallgrass savanna) on habitats of deeper soil and fewer rocks the natural range plant communities are generally more mesic. This more moisture-requiring glade vegetation is often ephemeral being limited mostly to the wetter, cooler season prior to onset of summer heat and the drier soil conditions. One of the more spectacular of these ephemeral glade range communities are in shallow depression on higher-clay content soils. These wet-season depression communities can be regarded as vernal pools, although that designations is not commonly applied to such range vegetation.

One of the more dramatic of these vernal pool communities (at least in wet, favorable years when its "in all its glory") is the one dominated by widow's cross (Sedum pulchellum). This annual stonecrop, with its water-efficient Crassulacean Acid Metabolism photosynthetic pathway, has a typical boom-or-bust population cycle. When this dainty forb makes a crop it makes one hellacious, one "helluva" crop. Such a crop of widow's cross at peak standing crop was presented in these two "photoplots".

Associated with widow's cross in the range plant community seen here was plains tickseed or calliopsis (Coreopsis tinctoria) and eastern pricklypear cactus with a few chert stones covered by various lichen species (These lichens were treated below.) Whereas the rock outrcrops immediately upsloep from theser vernal pools had lanceleaf tickseed (Coreopsis lanceolata), a perennial composite, as a locally dominant forb, the vernal pool community had the annual species C. tinctoria. Astute students already should have " picked up" the fact that eastern prickly (like all the Opuntia species) also have the Crassulacean Acid Metabolism photosynthetic pathway.

The annual life cycle and the Crassulacean Acid Metabolism pathway are plant adaptations (evolutionary, natural selection traits) to harsh, especially water-stressful, edaphic conditions. Annuals with their tough, durable, long-lived seeds are the textbook example of the therophyte life or growth form (Allaby, 1998).

These vernal pool communities were located just above the Ozark Plateau stream of Shoal Creek.

20. The ones below the "rock center"- Two close-in "photoquadrants" of the range plant community of a vernal pool that developed on small land depressions forming on high-clay soil at the base of chert glades in the Ozark Plateau of southwestern Missouri. There were some small chert rocks (such as the one in the center of the second "photoquadrant") these were not the large flagstone forms of chert outcropping that characterized the more typical chert glades located upslope from the lower, deeper (less shallow) soils.

The overwhelming dominant of vernal (seasonal aspect) plant society (of the climax range vegetation) was widow's cross, an annual stonecrop. Widow's cross was in full-bloom to early fruit phenological stage so as to be at peak standing crop. Eastern pricklypear cactus (center of first slide) and plains tickseed or calliopsis, an annual composite of the Heliantheae tribe, were distant associate species to widow's cross.

The small chert fragments (larger stones to small rocks) were the habitat of various species of lichens and mosses. Details of the range vegetation--includes thallophytes and bryophytes--of chert glade vernal pools were featured below.

Location note: slides of plant species found in the above and below images of vegetation were presented at the end of this subsection.

21. Below the big rocks- The range vegetation that developed on the outer margins of vernal pools which formed on depressions in high-clay content soils at the lower elevational end of chert glades. The climax vegetation in the depressions was presented in the immediately preceding slide-caption set. These vernal pool plant communities were overwhelming dominated by the annual stonecrop, widow's cross. On the perimeter or margins of these depressions the glade vegetation also had widow's cross as a major to the dominant species, but this foliar cover of widow's cross was not nearly as overwhelming as farther into the depressions. This condition was shown in the first of these two slides.

The second slide presented a greater area of coverage (so widow's cross was less conspicuous) which showed that there was proportionately greater cover of one of the associated species, plains tickseed or calliopsis (an annual composite). Also featured in this second slide was was a low "rock pile" (center of photograph). The surfaces of the chert stones in this a natural and somewhat circular arrangement were covered by various lichen species with mosses, widow's cross, Nuttall's yellow stonecrop, and starved or dwarf rosette panicgrass growing in spaces of soil within the pattern of chert fragments.

Details of this range vegetation within the circle of chert fragments was given in the next slide/caption set. Next slide, please.

22. On the smaller or outer rocks (in the inner circle)- Set of three progrsssively closer-in views (set of three "nested photoquadrants") of the range vegetation on and around small chert fragments in a depression in high-clay content soils at lower elevations of a chert glade in the western Ozark Plateau. These depressions functioned as what have been described as "vernal pools", localized natural pools or small bodies of ponded water forming during the wetter, cooler parts of spring through early summer. The general view of these depressions was presented and described above, including the range vegetation that developed on a small chert fragments naturally arranged in a circular pattern shown in the immediately preceding two-slide/caption set.

The first of these three slides was a top-down view of some of the lichen-covered chert stones. The principal lichen on these chert fragments was--depending on the rock chosen to concentrate on--either the foliose lichen, Cumberland xanthoparmelia lichen, or the cup lichen known as olive cup lichen or olive cladonia (Cladonia strepsilis). Seen in this first slide were plants of both Sedum species growing in these vernal pool and adjacent habitats: 1) widow's cross (S. pulchellum) and 2) Nuttall's yellow (or golden) stondcrop (S. nuttallii). Also present though not visible at this size was starved or dwarf rosette panicgrass.

The second slide presented both Nuttall's yellow stonecrop and widow's cross along with Cumberland xanthoparmelia lichen to their right. The third slide provideded a closer-up view of widow's cross and olive cup lichen in lower left corner plus a shoot of starved or dwarf rosette panicgrass, the principal grass of these vernal pool communities.

23. On the chert surface- Lichens' turn. Chert fragments ranging in size from small stones (about large pebble-size) to medium rock-size served as the habitat for thee species of lichen: 1) a foliose lichen, Cumberland xanthoparmelia lichen (Xanthoparmelia cumberlandia), 2) a cup lichen, olive cup lichen or olive cladonia (Cladonia strepsilis) and 3) a stippleback lichen common or silverskin stippleback (Dermatocarpon miniatum). This assemblage was presnted in the first slide. (A small plant of widow's cross snuck in this group shotin lower left corner.)

The second slide showed students a closer-in view of Cumberland xanthoparmelia lichen and just below it the blackish green of a byrum moss (Bryum pseudotriquetrum).

24. Where there's more water than rocks- A vernal pool that formed in a slightly greater depression on deeper, clayey, less stoney soil at the lower elevational end of chert glades in the Springfield Plateau. This vernal pool vegettion was dominated by Texas or spreading chervil (Chaerophyllum texanum) of the carrot or parsley family (Umbelliferae) and taper-tip flate-sedge (Cyperus acuminatus) of the Cyperaceae. The associate plant species of this deep vernal pool vegetation was Engelmann's spike-rush or spike-sedge(Eleocharis engelmannii), an annual member of Cyperaceae. Engelmann's spike-sedge was the sole plant species that grew in the very bottom or lowest point of the vernal pool. A fourth and the least common plant species of this deep vernal pool habitat was needle-point or two-flower rush (Juncus marginatus).

The first of these two slides presented a side-view of the overall range plant community of the deep vernal pool environment. This range vegetation was comprised of Texas chervil, an annual, and taper-tip flatsedge, a cespitose perennial. The second slide was a closer-in and topdown view of taper-tip flatsedge, which was more conspicuous, and Texas chervil (more cover in left side of this slide).

25. Winter dominant- Hairy lip-fern (Cheilanthes lanosa) growing on rock outcrops in a chert glade in the western Ozark (Springfield) Plateau in late spring. This pteridophyte is a locally dominant range forb in the spring to summer seasons (spring and summer societies of the climax vegetation and it is an overall dominant of the winter society. (See hairy lip-fern below for winter growth and habit.)

Wildcat Glade Natural Area, Newton County, Missouri. Early January; peak standing crop, mature spore phenological stage.

26. Little lips in the making- Mature fronds of hairy lip-fern with spore-bearing fronds. Adaxial (upper) surface and overall view in first slide and abaxial (lower or beneath) surface and closer-in view of fronds in second slide. The spore-bearing organs are on the underside (abaxial surface) of fronds and are called sori (plural; singular is sorus). Any standard Botany text will provide a good discussion of sexual reproduction in pteridophytes. Some of the older textbooks are better for giving more detail (in something besides Molecular Biology) and having line drawings that are superior to the current colored illustrations. Recommended Botany texts included Holman and Robbins (1939, ps. 506-508, 514-516), Cronquist (1971; ps. 371), and Wilson et al. (1971, ps. 611-619).

Wildcat Glade Natural Area, Newton County, Missouri. Early January; early frond growth phenological stage.

27. In winter's cold chill- Hairy lip-fern in its winter "paleage" growing on a rock outcrop in a chert glade in the Springfield Plateau. Last year's dead fronds (fern leaves) served as background for this year's new fronds which are extremely winter hardy. (These slides were taken on a frosty January morning with temperature about 12 degrees Fahrenheit.)

Hairy lip-fern is a major range plant species on Ozark Plateau glades. This pteridophyte is a dominant of the winter society of this climax vegetation.

Depending on taxonomist serving as the authority, hairy lip-fern has traditionally been included in the Polypodiaceae, the true fern family.

The third slide was improperly scanned (too much light--bleached out--and could not be corrected back to original image with Adobe Photoshop) with a Epson Perfection 600 scanner. Fortunately, the photographer's Nikon FM2 functioned perfectly on this cold winter morning. (It may well be that a poor craftsman blames his tools, but try getting shots like this with a pin-hole camera. Or, compare the performance of a quality brand like Nikon to inferion machinery like an Epson scanner.)

Back to the subject at hand, hairy lip-fern is well-adapted to the shallow, xeric, infertile environment of chert glades. While many fern species are better fit to live in moist environments hairy lip-fern makes its home on the harsh habitat of Ozark chert glades.

Wildcat Glade Natural Area, Newton County, Missouri. Early January; early frond growth phenological stage.

Speciation on Ozark (Springfield)Plateau chert glades- On the chert glades and habitats contiguous with (and in very close proximity to) the chert glade range site tbeing treated in this section there werethree Coreopsis species growing on four obviously different niches (one species per niche as per Gauss's Competitive Exclusion Principle) and physically inhabiting four different edaphic habitats. In addition to differences in edaphic features there were pronounced differences in timing of blooming with one ot the three species blooming one month later thatn the other two (late spring versus early summer) and, perhaps, most telling of all in life cycle of species with two species perennial and one an annual.

This was not quite up to Darwin's finches or tortoises, but it certainly was a textbook example of speciation and niche creation- occupation. Good source for teachers wishing to use less exotic and closer-to-home examples of natural selection.

28. On chert outcrops- Lanceleaf tickseed (Coreopsis lanceolata), a perennial and more-or-less cool-season species. Lanceleaf tickseed often grows as large stands (populations yet of clonal genotypes) over chert outcrops of extremely shallow soils. This is early spring with most blooming completed before hotter summer temperatures evaporate the limited water from the harsh edaphic environment. In wet springs it is not uncommon to see whole hillsides of chert glades covered with the conspicuous flowers of this species. It is a climax dominant of chert glades grasslands.

Chert rock outcrop, Wildcat Glade Natural Area, Newton County, Missouri. Late May- peak-bloom phenological stage.

29. Shoots on chert outcrops- Shoot features of lanceleaf tickseed showing the namesake lanceolate leaves that clasp the slender stem of this showy perennial composite. Lanceleaf tickseed is the dominant forb of the spring flora of chert glades, often the dominant range plant species of vernal societies.

Chert rock outcrop, Wildcat Glade Natural Area, Newton County, Missouri. Late May- peak-bloom phenological stage.

30. Heads on the chert outcrop- Capitula of lanceleaf tickseed showing characteristic outline of ray petals.

Chert rock outcrop, Wildcat Glade Natural Area, Newton County, Missouri. Late May- peak-bloom phenological stage.

31. Off to a deeper-soiled side- Details of shoot details (first slide) and of capitulum (second slide) of bigflower tickseed (Coreopsisgrandiflora). This species does not thrive on chert glades, but instead has its principal habitat more mesic environments with deeper soils. It does grow on edges of chert glades often in plant communities with more tree cover. It tolerates partial shade well.

Edge of stunted oak stand, Wildcat Glade Natural Area, Newton County, Missouri. Late May- peak-bloom phenological stage.

32. On wet glade soil with some depth- Dense population of plains tickseed or calliopsis (Coreopsis tinctoria) on a vernal pool on Ozark (Springfield) chert glades. This is an annual species with pronounced "boom and bust" years. Guess which this was. At base slopes or flatter areas around perimeters of chert glades where runoff and ponded water from heavy spring rains accumulates plains tickseed or calliopsis forms expansive populations that "are a beauty to behold".

In this habitat calliopsis was "accompanied by" the annual stonecrop known as widow's cross (Sedum pusillum) and grasslike plants.

Palatability of this species was unknown to the author, but plains tickseed is basically a typical annual forb adapted to disturbance and short-lived abundance of resources, namedly water on this habitat or localized range site.

Vernal pool, Wildcat Glade Natural Area, Newton County, Missouri. Early July; peak-bloom phenological stage.

33. Shoots on the wet glades- Details of shoot of plains tickseed, the dominant range plant on vernal pools on the adjacent "outskirts" of chert glades in the Springfield Plateau.

Vernal pool, Wildcat Glade Natural Area, Newton County, Missouri. Early July; peak-bloom phenological stage.

34. Tincture on wet rock piles- Capitula (heads) of plains tickseed or calliopsis growing on a vernal pool at base of chert glades in the western Ozark (Springfield) Plateau. Calliopsis was dominant to do-cominant with widow's cross on vernal pools.

Vernal pool, Wildcat Glade Natural Area, Newton County, Missouri. Early July; peak-bloom phenological stage.

35. Widowed in a vernal pool- Local dene stand of the annual plant called widow's cross (Sedum pulchellum) growing on a vernal pool within the overall chert glade range plant community in the western Ozark Plateau. There several Sedum species in North America almost all of which grow on harsh sites and microsites usually characterized as being rocky and/or with shallow soils. These habitats for the various Sedum species range from alpine to low elevation environments. (Several Sedum species were presented throughout Range Types Of North America; and, again, most of these were calling otherwise hostile habitats their home.)

Some Sedum species are perennials while others, such as widow's cross, are annuals. The annual species exhibit typical boom-and-bust, feast-or-famine yearly population fluctuations. The large dense stand on a vernal pool that was presented here grew in a year or, more specifically, spring-to-autumn growing season that was a Severe Drought (Palmer Index), but for a brief period there was heavy rainfall that flooded the vernal pool topography that permitted the extensive heavy population of widow's cross that served as the example for the chert glade range type. This was an instance of where the annual life cycle allocation of resources enabled this diminutive species to thrive when and where larger perennials were suffered to produce lesser amounts of biomass and, more importantly, lower levels of reproductive output.

Vernal pool, Wildcat Glade Natural Area, Newton County, Missouri. Early July; peak-bloom phenological stage.

36. Cross on the chert glades- Inflorescences of widow's cross in a spring pool at lower end of an Ozark (springfield) Plateau chert glade--and what beauties they are! Although these beauties grew in an overall drought year (officially Severe Drought throughout most of the growing season) short-lived heavy rain showers over the vernal pool relief produced the dense-population stands and the extremely large, gorgeous inflorescences in this one otherwise poor plant-producing spring.

Vernal pool, Wildcat Glade Natural Area, Newton County, Missouri. Early July; peak-bloom phenological stage.

37. Others of the vernal pools- Engelmann's spikerush (Eleocharis engelmannii) growing on an ephemeral (short-lived) wet depression that frequently fills with water during spring (hence, vernal pool). This species grew in the centermost (thus, most mesic or, temporarily, most hydric) part of vernal pools while widow's cross and plains tickseed grew on more outer (less mesic or hydric) parts and perimeter of these vernal pools.

These three species are annuals showing the survival value ("fitness" in terms of natural selection) of the annual growth or life form known as therophyte. Survival by means of sexual diospores (= propagules, germules, dissimules) is the ultimate plant adaptation to harsh or erratic environments such as ephemerally wet or moist habitats like vernal pools.

Engelmann's spikerush was growing in the vernal pool with tapertip flat-sedge or umbrella-sedge (Cyperus acuminatus) and needle-point or two-flower sedge (Juncus confusus).

Vernal pool, Wildcat Glade Natural Area, Newton County, Missouri. Mid-July; immature-fruit phenological stage.

38. Spikes on spikerush- Inflorescences of the spike type on shoots of Engelmann's spikerush growing on a vernal pool within an area of chert glades in the western Ozark Plateau.

Vernal pool, Wildcat Glade Natural Area, Newton County, Missouri. Mid-July; immature-fruit phenological stage.

39. Flat-out in vernal pools- Tapertip flat-sedge or tapertip umbrella-sedge (Cyperus acuminatus) growing in a vernal pool in or around chert glades in the western Ozark (Springfield) Plateau. There are numerous Cyperus species in the Ozark Plateau. Most of these grasslike species are adapted to moist habitats.

On this very mesic to hydric (at least temporarily) plants of tapetip flat-sedge were growing in the centermost (most hydric) portions of vernal pools accompanied by Engelmann's spikerush. Unlike spikerush (and forbs growing on the perimeter of vernal pools) tapertip umberella-sedge is a perennial. Like the annuals, however, tapertip flat-sedge exercised its option of sexual reproduction (as well as asexual or vegetative reproduction).

Tapertip flat-sedge was growing with Engelmann's spikerush and neeedle-point or two-flower rush (Juncus marginatus).

Vernal pool, Wildcat Glade Natural Area, Newton County, Missouri. Mid-July; ripening-fruit phenological stage.

40. Sex in the vernal mud- Spikelet-dense inflorescences of tapertip flat-sedge produced on a vernal pool in or near chert glades in the Springfield Plateau of southwestern Missouri. This perennial grasslike species used both sexual reproduction (seed produced in achenes, the dry fruit type of Cyperus species) and asexual reproduction via tillers.

Palatability (or lack thereof) or forage value for Eleocharis and Cyperus species growing in these vernal pools was unknown to this author.

Vernal pool, Wildcat Glade Natural Area, Newton County, Missouri. Mid-July; ripening-fruit phenological stage.

41. Needle-pointed in a pool- Single plant of needle-point or two-flower rush (Juncus marginatus) growing in a drying vernal pool at edge of chert glaces in the Springfield Plateau of southwest Missouri. This plant was "accompanied by tapertip flat- or umbrella-sedge and Engelmann's spikerush."

Vernal pool, Wildcat Glade Natural Area, Newton County, Missouri. Mid-July; ripening-fruit and near-dormancy phenological stage.

42. Flowered and pointed in a pool- Flower cluster (panicle) of needle-point or two-flower rush (first slide) and closer-in view of units of fruit (second slide) growing in a drying vernal pool at edge of chert glades. The fruit of Juncus species is a locucidal capsule.

Vernal pool, Wildcat Glade Natural Area, Newton County, Missouri. Mid-July; ripening-fruit phenological stage.

43. A Texan on an Ozark glade- Several annual plants (first or vertical slide) and part of the umbel inflorescence (second or horizontal slide) of Texas chervil (Chaerophyllum texanum) growing in a vernal pool at edge of a chert glade. Texas chervil grew in local "pure" (homogeneous) populations or single-species stands in association with the three grasslike species of tapertip flat-sedge or tapertip umberella-sedge, needle-point or two-flower rush, and Engelmann's spikerush plus the forbs widow's cross and plains tickseed or calliopsis.

Texas chervil is in family Umbelliferae. Like the species with which it was associated, this umberlliferous annual forb preferred wet habitats. In dry years Texas chervil was absent from the chert glade landscape.

Newton County, Missouri. Mid-July; late-bloom phenological stage.

44. A defining dominant- Two views of a large--though severely droought-stressed--plant of switchgrass, an upland ecotype. This plant was growing on a chert glade in the western Ozark (Springfield) Plateau in the second consecutive summer of Extreme Drought (Palmer Index). Switchgrass is one of the defining species of tallgrass prairie (Weaver and Clements, 1938, p. 518).

This example showed prairie dropseed in the early sexually reproductive stage (only two of the sexual shoots had emerged panicles). At this stage of phenology the long shoots (vegetative shoots that elongate relatively early in their growing season) had progressed to that part of the sexual stage when internodes elongate prior to emergence of panicles.

Wildcat Glade, Newton County, Missouri. Late July; peak phytomass, shoot elongation and panicle expansion stage of phenology.

45. Shoots of a seemingly unusual defining dominant- Two views into the interior of a severely drought-stressed plant of switchgrass growing on a xeric chert glade in the Ozark Plateau. These views presented details of culm and leaf. These organs were on the same plant seen in the two immediately preceding slides.

Prairiemen would not ordinarily expect to find (let alone actually find) switchgrass, the most mesic of the Four Horsemen Prairie Grasses (big bluestem, Indiangrass, and little bluestem being the other three) on this shallow, rocky, droughty range site, but such is the "fickleness of Mother Nature".

Wildcat Glade, Newton County, Missouri. Late July; peak phytomass, panicle expansion stage of phenology.

46. Leafy feature of an unexpected dominant- Leaves of an upland ecotype of switchgrass growing on a chert glade in th western Ozark Plateau. These drought-stressed leaves were on the plant presented in the two immediately preceding slide-caption sets.

Wildcat Glade, Newton County, Missouri. Late July; peak phytomass, full leaf expression.

47. Stressed sex of a rocky habitat dominant- Newly emerged panicle of switchgrass on shallow, rocky (chert) soil in the second consecutive summer of Extreme Drought. Panicles of switchgrass are typically large clusters of flowers--and in a favorable growing season--yield of caryopses. The just-exerted panicle seen in these two views had not had adequate time to expand to the degree it ultimately might have. Impact of drought stress was unknown (far from ideal conditions for plant growth or photography). The leaf immediately below (at peduncle of) panicle was the flag leaf.

Wildcat Glade, Newton County, Missouri. Late July; pre-anthesis stage of phenology.

48. Spikelets on a prairie panicle- Upper branches of the panicle of an upland ecotype of switchgrass with immature spikelets. Spikelets of the Panicum species (in fact, Paniceae tribe) typically have only a single, perfect terminal floret with a sterile floret subtending this fertile floret (Hitchcock and Chase, 1950, p. 24). The fruit is the "standard" caryopsis or grain.

Wildcat Glade, Newton County, Missouri. Late July; pre-anthesis stage of phenology.

49. Starved on a chert glade- Starved rosette panicgrass (Panicum depauperatum; subgenus, Dichanthelium) on a chert glade in the western Ozark (Springfield) Plateau. In spite of the designation referring to depauperate (stunted, starved, dwarfed; a runt, dwarf, etc) this species can grow to "normal" sizes for the rosette panicgrasses. This particular plant did, indeed, attain large size by standards for members of the Dichanthelium genus.

In recent decades numerous agrostologists have elevated subgenera, sections, and other subgeneric levels of Hitchcock and Chase (1950) to their own genera. It has been emphasized throughout this publication that the current author (a crotchy curmudgeon perhaps) has rejected most of this "taxonomic foolishness", including cladistics as a basis for biotic taxonomy.

Newton County, Missouri. Early July; vernal phase of annual growth cycle.

50. Another perspective- Side view of starved rosette panicgrass growing on a chert glade in the Springfield Plateau.

Newton County, Missouri. Early July; vernal phase of annual growth cycle.

51. Base on chert glades- Base of plant and basal leaves of starved roettte panicgrass on a chert glade in the Ozark (Springfield) Plateau.

Newton County, Missouri. Early July.

Chert Glade in Winter

The grassland aspect (the herbaceous component or, most specifically, the vascular plant somponent) of chert glades undergoes the same seasonal changes as those of surrounding tallgrass prairie and the herbaceous element of tallgrass-oak-hickory savanna. It is the much greater proportion of the thallophyte (primarily lichen) and bryophyte (moss) component of chert vegetation that prevails in winter (acutally through early spring as well). This nonvascular plant part of chert vegetation is primarily restricted to chert rock outcroppings and the intraspeces within the larger rock outcrops.

The following sub-section dealt with the winter vegetation society of the chert range type.

52. Gladed into winter- Three progressively smaller-scale views (three nested"photoplots") of an Ozark Plateau chert glade showing the winter aspect of this range type. Most of the brown- or tan-colored clumps are switchgrass the overall dominant plant species of this cover type. Other grasses included poverty dropseed (Sporobolus vaginiflorus), puffsheath dropseed (S. neglectus), little bluestem, big bluestem, winter bentgrass (Agrostis hyemalis), poverty or starved rosette panicgrass, and Canada wildrye. Eastern pricklypear cactus was the only woody plant present.

The chert outcrops were large to massive and covered a high proportion of the land surface. Outcroppings were mostly of the flagstone configuration (shape) often with prominent or conspicuous spaces of soil within them which showed impacts of geologic weathering. These cracks or spaces within flagstone arrangements were populated by trachyophytes (vascular plants), both pteridophytes like hairy lip-ferns and angiosperms including a diversity of both cool-season and warm-season forbs and grasses.

The surfaces of chert were largely covered by various lichen and moss species. The most abundant (having greatest cover) lichen was the foliose lichen, Cumberland xanthoparmelia lichen (Xanthoparmelia cumberlandia). Other lichens included olive cup lichen or olive cladonia (Cladonia strepsilis), and common or silverskin stippleback lichen (Dermatocarpon miniatum).

These views were immediately above Shoal Creek.

Wildcat Glade Natural Area, Newton County, Missouri. Early January. FRES No. 39 (Prairie Ecosystem), K-66 (Bluestem Prairie). SRM 803 (Missouri Glades). Closest biotic community unit in Brown et al. (1998, p. ) was Cold Temperate Grassland 142, Plains Grassland 142.1, Bluestem "Tall-Grass" Series 142.11. Range site is Limestone Rock Outcrop (Soil Conservation Service, 1989 did not list range sites for Newton County, Missouri) or Very Shallow with limestone flagstones for adjoining Ottawa County, Oklahoma (Soil Conservation Service, 1964). Ozark Highlands- Springfield Plateau ecoregion 39a (Chapman et al., 2002).

53. Dormancy (for some plants) on the chert glades- The winter society of climax range vegetation on an Ozark Plateau chert glade. Most herbaceous cover was switchgrass, the dominant vascular plant of this "rock garden" range. Other grasses included puffsheath dropseed, poverty dropseed, Canada wildrye, little bluestem, starved or dwarf rosette panicgrass, and some big bluestem which was a local dominant. The The surfaces of the chert outcrops were mostly covered by lichens especially Cumberland xanthoparmelia lichen and with less coverage by olive cup lichen or olive cladonia.

54. Thriving on a wet and cold glade- Yes, it even looks inhospitable, but this a prime season of growth for thallophytes on an Ozark Plateau chert glade. The temperature was 23 degrees Fahrenheit and there was a heavy overcast when your professor took these photographs, but various species of lichens and mosses were having a heyday. The dominant prairie grasses like the plant of big bluestem at right edge of the chert outcrop in center of the first slide were in dormancy while the annual grasses like puffsheath dropseed in upper center had completed their life cycle and were represented by their living, overwintering grains. There was also some poverty dropseed and several unidentifiable forbs.

Lichens, by contrast, were swollen and turgin with water that was not evaporating very fast in the cold, wet conditions of this early winter environment. The lichen with greatest cover on the chert surfaces was most Cumberland xanthoparmelia lichen, but the round, gray-green "puffball"-looking objects were lichens of olive cup lichen (Cladonia strepilis).

55. Who said we're "dormant"? Two "nested photoplots" of winter plant life on a chert glade in the western Ozark (Springfield) Plateau. Dead herbage in the first slide was that of tallgrass prairie grasses of which switchgrass was the dominant. There was also substantial dead biomass (specifically necromass) of big bluestem. Much of this necromass was also dead shoots of last year's growth of hairy lip-fern. The second slide (the "nested sub-photoplot") featured some of these dead shoots of hairy lip-fern, but the stars of this local show were the new, green and growing shoots of the current season's herbage of hairy lip-fern.

Most of the covering of chert surfaces was that of the foliose lichen, Cumberland xanthoparmelia lichen.

56. Showing the cold- Two slides of plant life on an Ozark Plateau chert glade at start of what turned out to be a cold winter. The first slide featured chert rocks covered primarily by Cumberland xanthoparmelia lichen with olive cup cladonia lichen also well-represented. The two tan or reddish brown clumps in center foreground in this first slide were two dead plants of the annual grass, poverty dropseed. There was also a dead sexual shoot of big bluestem from a large clump of that species in the upper left corner.

The second slide featured a poor "suffering" specimen of eastern pricklypear cactus that was dehydrated and showing the effects of a cold winter. (It was 23 degrees Fahrenheit at time of photograph, but temperatures had been below zero numerous mornings). Pricklypear can take the dehydration, but cold can set it back quite a bit even though this native species is right at home on chert glades.

57. Winter enough and then some- The cryptogams and At time of photograph the temperature was 23 degrees Fahrenheit, but temperatures had been below zero frequently even though winter was less than a month old.

Wildcat Glade Natural Area, Newton County, Missouri. Early January.

58. The spectrum of embryophytes- Plant species ranging from lichens, bryophytes, to angiosperm hardwood species to the dominant monocots including switchgrass and big bluestem on the rock outcrop habitat of a chert glade in the western Springfield Plateau in southwest Misssouri. Embryophytes are those plants which produce embryoes from sexual reproductionand have life cycles involving alternation of generations.

Mosses on the chert boulder were green byrum moss (Byrum pseudotriquetrum) and dry rock moss or silver beard moss (Grimmia laevigata).

Wildcat Glade Natural Area, Newton County, Missouri. Early January.

59. Two common bryophytes- Silver beard or dry rock moss (darker green) and green byrum moss (the lighter or yellowish green clumps) in their characteristic winter "coat" of sporophytic shoots growing on an outcrop of chert. Chert glades in the Springfield Plateau in southwest Missouri. These were the two were the most common (abundant) bryophyets in the cryptomaic layer or component of the rock outcrop portion of an otherwise tallgrass prairie.

Wildcat Glade Natural Area, Newton County, Missouri. Early January.

60. At peak in winter- Chert rock-covering winter growth of green byrum moss on a chert glade in the western Springfield Plateau. The bryophytic foliage growing here was the dene development of sporophytic shoots. Winter is typically the time of this sexual reproducction in the mosses of the Ozark Highlands.

Dead leaves ranged from post oak to big bluestem.

Wildcat Glade Natural Area, Newton County, Missouri. Early January.

61. Young stuff- Developing sporophytes (specifically, sporophytic shoots) of common green byrum moss at home on an outcrop of chert on a chert glade in the western Ozark (Springfield) Plateau.

Wildcat Glade Natural Area, Newton County, Missouri. Early January.

62. Showing the cold also- Three "nested photoquadrants" (three slides of progressively closer camera distance) of thallophytic and bryophytic plant life on a chert gladee in the western Ozark Highlands in southwest Missouri. There were two species of lichen prominent in these "plots": 1) Cumberland xanthoparmelia lichen (Xanthoparmelia cumberlandia) and 2) olive cup lichen or olive cladonia (Cladonia strepilis). The bryophyte was common green byrum moss (Byrum pseudotriquetrum) seen as the more conspicuous green "balls". Shoots of big bluestem and switchgrass were lying across these non-vascular plant members of the chert glade climax vegetation.

These non-vascular members of the range plant community were filled "pert nar to busting" with water in the cold, low-rate-of-water evaporation atmosphere of an Ozarks early winter. Made for fine slide-shooting picture as long as numb hands could work a 35mm Nikon fully manual (cold does not phase a Nikon fm, at least not at temperatures that this rangeman endures).

63. Still showing cold- Some more "family portraits" of lichens on an Ozark Plateau chert glade in winter. Lichens thrive in moisture-rich air, a condidtion sometimes seen in a wet winter (especially when moisture is coupled with cold temperatures that reduce rates of evaporation). The first slide was a "group shot" of Cumberland xanthoparmelia lichen, olive cup lichen or olive cladonia (the most abundant species seen here), and cup lichen (Cladonia dimorphoclada), a fruticose lichen, which is the nice specimen in right fore- to midground.

The second slide had as its centerpiece a chert rock covered with Cumberland xanthoparmelia lichen with shoots of switchgrass and poverty dropseed surrounding the lichen-covered rock.

Often times lichens are larger and more prominent in winter than in summer due to less evaporation of water from lichen tissues. Shown immediately below were some examples of major species of lichens including a xanthoparmelia lichen or rock-shield lichen which is a foliose lichen,commonly known as the Cumbberland rock-shield lichen. (Xanthoparmelia cumberlandia) on the chert glade being presented herein..the Xrock-shield lichen.

64. On a rock and a hard place- Cumberland xanthoparmelia lichen (Xanthoparmelia cumberlandia), a foliose lichen on chert rock outcrop on chert glades, Springfield Plateau. Grass by rock outcrop in second slide was an upland, xeric ecotype of switchgrass (Panicum virgtum).

Wildcat Glade Natural Area, Newton County, Missouri. Early January.

65. Foliose flowing over rock- Cumberland xanthoparmelia lichen, a foliose lichen, covering outcroping of chert in an Ozark (Springfield Plateau) chert glade. Grasses in the first (upper) slide were switchgrass and poverty dropseed.

Wildcat Glade Natural Area, Newton County, Missouri. Early January.

66. Got 'ya covered- A progressively closer-in view of Cumberland xanthoparmelia lichen, a foliose lichen, sprawling over chert rock on an Ozark (Springfield Plateau) chert glade on a cold morning in early winter.

Wildcat Glade Natural Area, Newton County, Missouri. Early January.

67. Thallic details- Close up view of thallus of Cumberland xanthoparmelia lichen, a foliose lichen on chert rock. This lichen species is in family Parmeliaceae, order, Lecanorales.

Wildcat Glade Natural Area, Newton County, Missouri. Early January.

68. Nice family and friend- Group (upper or first slide) and "friend", a single plant (second or lower slide) of olive cup lichen or olive cladonia (Clatonia strepilis), a foliose lichen, on a chert glade in the western Ozark (Springfield) Plateau. The forbs growing among the olive cup lichen in the first slide could not be identified. The grass species were puffsheath dropseed with some poverty dropseed both of which are annual species.

Wildcat Glade Natural Area, Newton County, Missouri. Early January.

69. Up-close look- Cladonia strepilis, a foliose lichen, on a chert glade in the western Ozark (Springfield) Plateau. This interesting symbiotic creature is in family Cladoniaceae of the Lecanorales order. Cladonia species are in family Cladoniaceae of order Lecanorales. These specimens were water-filled, turgid thallus bodies. Wet, cool to cold atmospheres are favorable to lichens absorbing and retaining water so as to make nice photographs (as long as photographer's hands do not "freeze up" on them).

Wildcat Glade Natural Area, Newton County, Missouri. Early January.

70. Another representative- (Cladonia dimorphoclada), a fruticose lichen, growing on a chert glade in the western Ozark (Springfield) Plateau. Another member of family Cladoniaceae.

Wildcat Glade Natural Area, Newton County, Missouri. Early January.

71. Does this look silver to you? - Silverskin lichen (Dematocarpon miniatum) on a chert glade in the western Ozark (Springfield) Plateau. Epson Perfection 600 scanner really messed this one up, but even with the botched scanning job viewers can still get the idea that this lichen has a light chocolate color. This mutualistic critter is in family Verrucariaceae of the Verrucariales order.

Wildcat Glade Natural Area, Newton County, Missouri. Early January.

72. Lichens and moss on the rocks- Olive cup lichen or olive cladonia (Cladonia strepilis) and common green byrum moss (Byrum pseudotriquetrum) on a chert glade in the western Ozark (Springfield) Plateau. Lichens and mosses swell to considerable degree in moist, cool to cold weather such that their details can be presented to good advantage.

Wildcat Glade Natural Area, Newton County, Missouri. Early January.

73. Hard rock and soft moss- Common green byrum moss (Byrum pseudotriquetrum) growing on a chert glade in the western Ozark (Springfield) Plateau.

Wildcat Glade Natural Area, Newton County, Missouri. Early January.

74. Dry and on the rocks- Dry rock moss or silver beard moss (Grimmia laevigata) growing beside Cumberland xanthoparmelia lichen at left. These interesting plants were growing on a chert glade in the western Ozark (Springfield) Plateau in the dead of winter. Wait, who said we're dead in winter on a chert glade?

Wildcat Glade Natural Area, Newton County, Missouri. Early January.

Recommended references- A good field guide to the more common cryptogams (including lichens as well as bryophytes) of the chert glades was the delightful work of Thomas and Jackson, 85). Sumary coverage of glade plant communities was provided by Ware (2002).

Dolomite Glade

75. Ozark Bald Knob- This is a dolomite glade or dolomite barrens in the Ozark Plateau. Such distinctive habitats and their vegetation are known to the local hillsmen as "balds" or "bald knobs". In virgin condition dolomite balds were almost entirely grasslands devoid of woody growth except for scattered individuals of yellow-wood or smoke tree (Cotinus obovatus), the most common shrub, smooth sumac (Rhus glabra), skunkbush sumac, sassafras (Sassafras albidum) red bud, and persimmon (Diospyros virginiana), all of which are in this scene. Dominant plants are the prairie grasses Indiangrass, prairie dropseed, and big bluestem with sideoats grama, poverty dropseed (Sporobolus neglectus), and Junegrass (Koleria cristata) as associates. The dominant forb which is co-dominant with the three dominant grasses is prairie dock (Silphium terebinthinaceum). Forb associates include calamint (Satureja arkansana), longleaf bluets (Houstonia longifolia), and spike lobelia (Lobelia spicata). Except for the dominant prairie dock, composites are rare and locally limited to yellow coneflower (Echinacea paradoxa), gayfeather (Liatris spp.), and compassplant (Silphium laciniatum). Lichens were common only on xeric microsites and much less developed than on the chert glade seen immediately above. The tall blue-green grass in the right foreground is Indiangrass, the dominant species of the bald. The tall yellow-flowered composite in center foreground is prairie dock, the dominant forb.

The Ozarks are ancient mountains worn down to their roots. The dolomite geologic strata are of the Cambrian-Ordovician formations, some of the oldest in these "everlasting hills". Note the concentric ring arrangement of the Cambrian-Ordovician formations.

Note also that this is bald is relatively free of invading trees, which unfortunately surround this oasis of pristine vegetation. The dominant invaders are eastern red cedar (Juniperus virginiana) and Ashe cedar or Ashe juniper (J. ashei). This glade has been kept open by prescribed burning which handily controls the non-sprouting noxious cedars yet allows maintenance of endemic shrubs like yellow-wood. Several ecological reviews of Ozark vegetation prior to European settlement have been made based on surveyors records, travelers journals, Army accounts, etc. (Steyermark, 1940; Beilmann and Brenner, 1951; Howell and Kucera, 1956; Steyermark, 1959). All these ecological surveys proved conclusively that throughout the Ozark Mountain Region oak-hickory-pine forests had expanded and invaded the pre-white man prairies and savannas. Cessation of fire along with farming, overgrazing, and commercial activity weakened the prairie sward and allowed the woody invasion that reached proportions of massive afforestation. The Soil Survey of Taney County Missouri, the adjoining county to the west of this glade, cited a mean fire frequency of 3.2 years on some Ozark glades and credited this as "the main reason for this open grassy upland landscape" (Natural Resources Conservation Service, 1996, p. 41).

A more colorful testimony of the original openness of these balds is the gripping tale of the "Bald Knobbers", post-Civil War "night riders" (vigillantes) who held clandestined meetings and lite signal fires atop bald knobs so as to be able to spot any unwelcome visitors and send coded messages to citizen peace officers (at least before vigilante justice degenerated into blood feuds). Today, most of the Bald Knobbers grassland meeting places are forests or cedar thickets.

McClurg Glade, Ava Ranger District, Mark Twain National Forest, Ozark County, Missouri. Estival aspect, July. No specific FRES or Kuchler designations for vegetation at this restricted scale, but it is primarily an "island"of FRES No. 39 (Prairie Ecosystem), K-66 (Bluestem Prairie). However, at landscape scale this is K-74 (Cedar Glades, Quercus-Juniperus-Sporobolus) which is contradictory because by definition glades are open areas of herbaceous vegetation so there is no way that oak and juniper could be the dominant genera. SRM 803 (Missouri Glades). Ozark Highlands- White River Hills Ecoregion, 39c (Chapman et al., 2002).

76. Bald knob range- Two photo-plots of a dolomite glade in the Missouri Ozarks maintained (more-or-less) by prescribed fire consistent with the natural fire regime. In the first photograph eastern red cedar and Ashe or post cedar were reinvading the bald knob. This is the Kuchler (1964; in map of Garrison, 1977) physiogonomic type designated Cedar Glades which recognized Juniperus virginiana and J. ashei as the dominant woody species of this potential natural vegetation. In absence of fire (and probably aided by improper grazing, especially overgrazing) these two junipers increase to such cover and density to result in a post-climax juniper woodland instead of the climax juniper-tallgrass savanna.

In the second photograph junipers were largely limited to the rock outcrop perimeter of the dolomite glade (where there is insufficient fuel to carry fire that kills these non-sprouting conifers). Instead the invading woody species was smoke tree (smoke bush) or yellow-wood. Also on the perimeter were individuals of smooth sumac, skunkbush sumac, sassafras, and redbud. These shrubs are well-adapted to fire. In fact, smooth sumac typically increases with range burning. Yellow-wood matures quickly and becomes senescent, but fire serves to "rejuvenate" older individuals of smoke tree through resprouting as was case for the young stump sprout in left center foreground of this second photograph.

Major (dominant) grasses were Indiangrass, prairie dropseed, and big bluestem. Numerous individuals of Indiangrass were readily distinguishable in these photographs as photographs as the tallest, bluish-green clumps of grass. Interestingly and for unknown reasons, many of the grass clumps of light-green (a "washed-out" green) color were also Indiangrass. In fact, several of these Indiangrass plants (prominent in center midground of first slide) had shoots of both the regular chalky blue coloration and the uncommon pale green color. Indiangrass is a long-shoot tallgrass that elongates the culm early in the warm growing season whereas big bluestem is a short-shoot tallgrass species that does not elongate the culm until much latter in the growing season (often two months or more after Indiangrass). Large specimens of Indiangrass were visible in front of the yellow-wood sprout in the second slide. Dead shoots topped with prominent panicles were specimens of the cool-season (hence, now dormant) Junegrass.

A nice specimen of prairie dock, the common dominant forb on bald knob glades, was in front of the young cedar in center foreground of the first slide. White-tailed deer (Odcoileus virginianus) bucks had been burnishing their antlers on this sapling juniper resulting in severe damage to the conifer and range improvement (aid in restoring the range plant community to potential natural vegetation) on this dolomite glade.

McClurg Glade, Ava Ranger District, Mark Twain National Forest, Ozark County, Missouri. FRES No. 39 (Prairie Ecosystem), K-66 (Bluestem Prairie) and K-74 (Cedar Glades). Prescribed fire has been used here at intervals of three to eight years. SRM 803 (Missouri Glades). Ozark Highlands- White River Hills Ecoregion, 39c (Chapman et al., 2002).

72. Grass and dock on the glade- Details of species composition of the herbaceous layers of this Ozark glade range. This example of bald knob vegetation was a consociation of prairie dropseed. It was a example of Clements' true prairie (in contrast to tallgrass prairie dominated by bluestems and Indiangrass). Tallgrass species of Indiangrass, big bluestem, and little bluestem (in that order) and prairie dock were associates to prairie dropseed. Other grasses included sideoats grama, poverty dropseed, and Junegrass. Other forbs were gayfeather willow-leaf sunflower (Helianthus salicifolius), compassplant, calamint and longleaf bluets.

The sward of this bald knob range had been maintained by three prescribed fires in recent records: 6, 12, and 22 years before time of this photograph.

FRES No. 39 (Prairie Ecosystem), K-66 (Bluestem Prairie) and K-74 (Cedar Glades). Prescribed fire has been used here at intervals of three to eight years. SRM 803 (Missouri Glades). Ozark Highlands- White River Hills Ecoregion, 39c (Chapman et al., 2002).

73. Detail of the vegetation of McClurg Glade- Prairie dock and prairie dropseed are the dominants; Indiangrass is the local associate. McClurg Glade, Ava Ranger District, Mark Twain National Forest, Ozark County, Missouri. Estival aspect, July. FRES No. 39 (Prairie Ecosystem), K-66 (Bluestem Prairie) and K-74 (Cedar Glades). Prescribed fire has been used here at intervals of three to eight years. SRM 803 (Missouri Glades). Ozark Highlands- White River Hills Ecoregion, 39c (Chapman et al., 2002).

Not the most distinctive but the defining species- Several plants of prairie dropseed, part of a local consociation, (first slide) and a large plant of prairie dropseed with prairie dock, dominant forb of many Ozark glades, in background (second slide). These two "photo-quadrants" were taken along a rural road where the introduced pasture grass, tall fescue (Festuca arundinacea) had all but eliminated native chert grasslands on which prairie dropseed and prairie dock were conspicuous dominant species. A little remnant of a unique grassland now almost lost.

McDonal County, Missouri. late July; pre-bloom stage both species.

Featuring a defining range plant- Of all the native plants growing on the more eastern prairies and also on the true prairie association or cover type none elicits as many questions and provokes as much comment as does prairie dock (Silphium terebinbinthinaceum). The combination of large, robust plant size; immense basal leaves; and brillian; yellow heads at peak bloom makes for the most eye-catching and comment-evoking plant on the prairies and glades. Nothing would do but to devote the appropriate space to such a marvelous prairie forb.

74. Tough forb in drought- Prairie dock (Silphium terebinbinthinaceum) growing on a chert glade in the Springfield Plateau during the second consecutive year (second warm-growing season in a row) of Extreme Drought (Palmer Drought Index) ). Prairie dock is one of the most distinctive and frequently asked-about plants on chert and limestone glades in the Ozark Plateau. It is one of three major Silphium species on this general range type.

McDonald County, Missouri. Late July; prebloom (but flower buds died due to drought stress).

75. Better luck (more rain) next year- The same plant of prairie dock as shown in the immediately preceding slides except these leaves and stems were in the next year, which though still a drought year, was a somewhat wetter and much cooler (less evaporation of soil water) warm-growing season. This year was only Severe Drought (Palmer Index). Plant was growing on edge of a chert glade in the Springfield Plateau section of the Ozark Plateau physiographic province.

McDonald County, Missouri. Early August; full-bloom stage.

76. Big leaf on a tough prairie- Leaf of prairie dock growing at edge of a chert glade in the Ozark (Springfield) Plateau in southwestern Missouri. This leaf was one of the leaves presented in the immediately preceding slide though taken from a different angle (camera focal point/position).

McDonald County, Missouri. Early August; full-bloom stage.

77. High beauty on a chert glade- Inflorescence (first of four slides) and one of the four capitula (heads) in the top slide shown separately (next two slides) of prairie dock. This was the inflorescence and heads atop the basal leaves and shoots of the same plant proudly displayed above. The fourth slide was of another head produced on this same plant in the succeeding year.

McDonald County, Missouri. Early August; full-bloom stage (no doubt of that).

78. Longleaf bluets (Houstonia longifolia)- A common forb on the more xeric microsites of dolomite glades or bald knobs. McClurg Glade, Ava Ranger District, Mark Twain National Forest, Ozark County, Missouri.

79. Purple bald knobber- Purple Cobey's beard-tongue (Penstemon cobaea var. purpureus), an endemic to the limestone glades (bald knobs) of the White River portion, Springfield Plateau of the Ozark Highlands province (Steyermark, 1963, p. 1343). This gorgeous taxon has become a favorite among native plant enthusiasts for growing (from seed) in native landscapes.

Ozark County, Missouri. Late May; peak-bloom phenological stage.

80. Flowers of a purple bald knobber- Flower cluster (first slide) and close-in views of two individual flowers of purple Cobey's beardtongue on a glade or bald knob in the White River portion of the western (Springfield) Plateau. Not only is this endemic prairie forb almost unbelieveably beautiful, it is also highly palatable to grazing animals. Given its palatability and endenism, the purple-flowered variety of Cobey's penstemon (beardtongue) is relatively scarce.

The color of the two flowers (second slide) was not accurately or correctly scanned (by an Epson Perfection 700 scanner). The purple of the flowers in the first slide in this slide-caption set and that of the same plant in the immediately preceding slide of this taxon was correctly reproduced or transmitted by this same scanner.

Ozark County, Missouri. Late May; peak-bloom phenological stage.

81. Invasion of dolomite glades or bald knobs by eastern red cedar, primarily, and with some Ashe cedar. At landscape scale this vegetation was mapped by Kuchler (1966) as K-74 (Cedar Glades, Quercus-Juniperus-Sporobolus) which as shown above was contradictory because grassland is climax and oaks and junipers are at best only aspect dominants. True dominants are Indiangrass, prairie dropseed, and big bluestem. Detailed studies clearly proved that woody invasion occurred and caused (or is the result of) range retrogression due mostly as a result of fire suppression. Fortunately that dim-wit bruin Smokey Bear died (cremation would have been more appropriate than his expensive burial). Now the Forest Service is using prescribed fires to clean up the bald knob country and restore the climax prairie vegetation. It was none-too-soon as seen on these cedar-invested glades over which a fire tower stands sentinel. How ironic, prophetic, and symbolic of good intentions that wrought bad! Fortunately man is a thinking animal.

Three Sisters viewed from Gladetop Trail, Caney Campground, Ava Ranger District, Mark Twain National Forest, Ozark County, Missouri. Estival aspect, July. FRES No. 39 (Prairie Ecosystem), K-74 (Cedar Glades). SRM 803 (Missouri Glades). Ozark Highlands- White River Hills Ecoregion, 39c (Chapman et al., 2002).

82. Cedar encroachment on Ozark Glades- Eastern red cedar and Ashe or post cedar with "tag-along" individuals of yellow-wood or smoke tree, sassafras, persimmon, smooth sumac (which increases rapidly and profusly with fire), and chittam-wood or gum-elastic (Bumelia languinosa var. albicans= B. languinosa var. oblongifolia).

This invading woody component was on the edge of the glade presented in the immediately preceding seven photographs where this glade joined unburnt portions. The woody vegetation at this edge served as a control plot from which response to prescribed fire of range vegetation on the prescription-burnt glade could be compared. Patchworks or mosaics of such burned and unburned range vegetation on Ozark Highlands handily fit the concept in Landscape Ecology and Wildlife Management of edge effect. Small tracts of cedar-dominated vegetation were patches within a matrix of glades just as at larger spatial scale glades were patches in an overall matrix of oak-hickory forest (patches inside patches).

Edges between adjoining dolomite glades and cedar-infested glades were small ecotnes that benefited wildlife species. However, the ultimate state of this Ozark Plateau range type in absence of natural fire or prescribed fire that simulated natural fire regimes is all too often a cedar (juniper) woodland or, ultimately, a cedar forest of closed canopy and no understorey, often with accelerated erosion of soil surface.

The sermon for this range type is a reversal of words, but the same moral as "repent or burn". On range such as this the prophetic message is "repent and burn or regret and perish".

83. The original Ozark hillsman- A beautiful male mountain boomer or collared lizard (Crotophytus collaris) at home in his kingdom on a dolomite glade commonly known as bald knobs to the hill folk. This is the State Reptile of Oklahoma and the original Ozark hillbilly. This defiant, fearless (the photograph was taken with a normal 50mm lense and the Bald Knobber's indulgence) little carnivore reminds the Range Management student that the trophic level structure of food chains in range ecosystems includes secondary or tertiary consumers that eat primary consumers (herbivores). As this Ozarker stands at the doorway of his dolomite castle he also reminds conservationists that the secret to wildlife abundance is habitat management. (The habitat component shown here is cover.) Ava Ranger District, Mark Twain National Forest, Ozark County, Missouri. July.

Granite Barrens

84. Complex of Stony Land- Rock outcrops are some of the most widely distributed forms of barrens. These occur on small areas or in scattered restricted locations within a landscape-scale or regional-sized general plant community (eg. rock outcrops on hill sides or mountain tops within a grassland association or forest formation (eg. granite outcrops within the Sierra Nevada Mixed Conifer Forest and in the Appalachian Highland or Ozark Plateau of the Eastern Deciduous Forest Formation).

Closely related to rock outcrops are talus or scree slopes, cliffs and bluffs, the various kinds of rock glades, sand and gravel bars along streams, sand dunes, and beach and dune habitats. These sometimes occur together and, as the case with the continuum of savanna vegetation, intergrade to the point that it becomes arbitrary to distinguish an outcrop from a rock glade from a talus slope. These all share the common feature of having immature, shallow soils, parent material on or close to the ground surface, and rapid rates of water runoff such that the habitats are droughty, generally harsh, and overall hostile to plant life. Nelson (1987, p. 99) designated these as primary natural communities which is certainly consistent with primary succession, primary materials, primary structure, or even primary schools.

The complex of primary areas and range sites seen here is in the Wichita Mountains of Oklahoma. Coming down from the top of this weathered mountain are the following designated categories of land: 1) Granite Outcrop Escarpment, 2) Granite Rock Outcrop, and 3) Stony Rock Land which in turn is divided into 4) Hilly Stony range site and 5) Hilly Stony Savanna range site. These were given in various geologic references and range site descriptions in Soil Conservation Service Soil Surveys (eg. Commanche County, Oklahoma). The vegetation is extremely variable and different distinct plant communities could be recognized at different spatial scales. Species include eastern red cedar, netleaf hackberry (Celtis reticulata), scrub form of blackjack and post oaks, slippery or red elm (Ulmus rubra), smooth and skunkbush sumac, big bluestem, Indiangrass, little bluestem, sideoats grama. Soils are mostly of the Lithosol great group (the 1938 soil classification system). It is all Land Capability Unit VIII.

Wichita Mountains National Wildlife Refuge, Commanche County, Oklahoma. Autumnal aspect, October. This is an "island" of rock outcrop or barrens within the "sea" of the mixed prairie range type (FRES No. 38 [Plains Grassland], K-62 [Bluestem-Grama Prairie]) There is no appropriate FRES or Kuchler Unit designation for this rock outcrop. As much as anything else it resembles a scrub form of Cross Timbers and legitimately could be interpreted as such (FRES No. 15 [Oak-Hickory], K- 75 [Cross Timbers]), but the author felt that it was unique from that (and all other vegetation) and should be categorized as a savannah or barrens. Central Great Plains- Wichita Mountains Eoregion, 27k (Woods et al., 2005).

85. Mountaintop savannah- Here on the top of Mount Scott (elevation 2464 feet) in the Wichita Mountains is a big bluestem-eastern red cedar savanna. Indiangrass and little bluestem are tallgrass associates and sideoats grama is the midgrass associate. The dominant forb is broom snakeweed (Gutierrezia sarothrae). Eastern red cedar is wind-pruned thereby having a semi-krumholtz form and forming a tallgrass-juniper savanna or perhaps a mixed prairie-juniper savanna. There is some Ashe juniper (Juniperus asheii) in the Wichitas, but eastern red cedar (J. virginiana) is overwhelmingly the dominant cedar. On dry, rocky hill tops like this the juniper would be less apt to burn and would logically be a component of the climax community such that the potential natural vegetation has to be interpreted as a savanna. As discussed under the previous slide no FRES or Kuchler Units exist at this restricted scale. Likewise there is no SRM rangeland cover type although some of the Texas Edwards Plateau types have a similar floristics and habitat (eg. SRM 733, Juniper-Oak Woodland). Consistent with the granite parent material of this "boulder field" and the inclusion of glades as a form of barrens and primary natural plant community it seemed that the most likely published classification of this unique range vegetation was Cedar Glades (Kuchler Unit 74) or, alternatively, some designation based on parent material (eg. Granite Glade or Granite Rock Outcrop) or perhaps a combination of geologic material and dominant species (eg. Granite Cedar Glades, Granitic Bluestem-Cedar Savanna). The range site designation is most fitting: Hilly Stony Savanna.

Wichita Mountains National Wildlife Refuge, Commanche County, Oklahoma. Autumnal aspect, October.Central Great Plains- Wichita Mountains Eoregion, 27k (Woods et al., 2005).

86. Granite Rock Outcrop- This boulder field resembles the boulder fields of the alpine ecosystem as do the wind-pruned near-krumholtz forms of eastern red cedar. Granite rock outcrop habitats and natural plant communities were discussed variously in Anderson et al. (1999), but none of this dealt with the exact kind of granite outcrop seen here. Big bluestem and Indiangrass are dominant grasses.Shrubs include scrub blackjack oak and Texas red oak or Spanish oak (Quercus shumardii var. texana) which is the larger shrub in the center skyline. Smooth and skunkbush sumac are visible in foreground and eastern red cedar is in far background. There is a diverse lichen community on the granite boulders. This plant community corrresponds closely to that of the chert glades which, as mentioned in the preceding slide, prompts designation of this as a granite glade. The tallgrass-oak-juniper-sumac savanna community is aptly designted as Hilly Stony Savanna range site. Wichita Mountains National Wildlife Refuge, Commanche County, Oklahoma. Autumnal aspect, October. Central Great Plains- Wichita Mountains Eoregion, 27k (Woods et al., 2005).

Serpentine Barrens

87. Serpentine Barrens Rock outcrop- An extremely harsh edaphic environment created by the presence of the mineral serpentine is one of the more common forms of barrens along the Pacific Coast. When serpentine barrens is combined with rock outcrop, one of the most common forms of barrens continent-wide, the habitat is definitely BAD.

Serpentine forms a unique, if harsh, habitat and, following an initial period of mostly being ignored by ecologists, serpentine sites became a "pet" subject of study in the Pacific Region. Readers are first referred to the relatively recent reviews by Kruckeberg (1984) and Kruckeberg in Anderson et al. (1999, ps. 309-321) as well as Tyndall and Hull in Anderson et al. (1999. Ps. 67-82). The older work, some of it classic, was given in the references of these recent reviews.

Serpentine is a mineral conisisting basically of a variable hydrous magnesium silicate, a complex magnesium-rich silicate- chiefly as Mg₃Si₂O₅(OH)₄- in fine waxy masses (but locally fibrous as asbestos), having colors ranging from dull or light green to dark green to black with intermediate brownish-red or yellow and often with a mottled appearance. Serpentine formed by alteration of peridotite (olivine) in presence of water and thus can occur in or as rock, both altered peridotite and pyroxenites. Serpentine is one of the ultramafic rocks (igneous rocks composed mostly of mafic minerals which are dark-colored iron-magnesian rock-forming silicates with olivine, chromite, and pyroxene as major minerals).

Serpentine parent material weathers into soils rich in magnesium, iron, nickel, and silicates. Serpentine soils have montmorillonite as the common clay mineral. Serpentine soils include both alluvial and upland series. The upland serpentine soils are azonal soils that are stony, shallow, and lack regular profile development. In addition, serpentine soils have high levels of exchangeable magnesium and exrtremely low levels of exchangeable calicum. They are typically deficient in nitrogen and phosphorus among macrominerals and molybdenum among trace elements while containing hyper-levels of the heavy metals iron and nickel. This creates a "serpentine syndrome" of infertile soils combined with toxicity of magnesium and heavy metals (Kruckeberg, 1984, ps. xii, 6-8, 18-26). In short, not the place for a plant to plan on having a party.

Yet, certain plants survive, if not thrive, here. Some of them are serpentine endemics. Others have distribution patterns varying between serpentine and non-serpentine soils while yet others are categorized as indifferent (= bodenvag= ubiquist) species and occur on and off serpentine soils with no apparent taxonomic differences within species on both categories of soil (but ecotypic variation cannot be ruled out definitely). See Kruckeberg (1984, ps. 34-44) for discussion of plant species relations on serpentine soils.

Response of range animals to serpentine sites and vegetation thereon has received much less study than has the more basic plant response.

The Serpentine Barrens Rock Outcrop Site seen here is dominated by big squirreltail (Sitanion jubatum) which is one of the indifferent or bodenvag serpentine species. This is a very restricted site (microsite) in the northern Coast Range. The surrounding oak woodland on either side and the California annual grassland behind are not on serpentine soils. Foster Pasture, University of California Hopland Field Station, Mendocino County, California. Late May, nearing end of growing season.

Serpentine Barrens were too small to be mapped at the scale used by Kuchler (1966, 1977), but this local site is obviously an "island" of the California bunchgrass prairie (K-41, California Steppe). As discussed earlier, SRM failed to designate and describe California steppe or Pacific bunchgrass prairie.

88. Serpentine Barrens range site of California annual grassland- Kruckeberg (1984, p. 41) listed a number of Eurasian annual grasses which he felt must be indifferent (= bodenvaq or ubiquist) species on serpentine and non-serpentine sites. He listed wild oats, soft brome, red brome, nitgrass (Gastridium ventricosum), and foxtail fescue (Festuca megalura). All of these species except wild oats are common on the parcel of naturalized annual grassland in this photograph. There is a relatively higher proportion of lower-seral stage species in this community with soft chess (= soft brome) being the only species that sometimes responds as a naturalized decreaser. Almost all of the grasses seen here are naturalized invaders like red brome, nitgrass, and foxtail fescue. Rosiere et al. (1986) studied the range plant community and domestic sheep diets on this same pasture and compared them to those from non-serpentine range sites on this same ranch. They found that there were few real differences in diets and the main difference in range species compostion was a lower proportion of decreaser and increaser grasses on the Serpentine Barrens.

Foster Pasture, University of California Hopland Field Station, Mendocino County, California. Late May. FRES No. 42 (Annual Grasslands Ecosystem), Kuchler (1966, 1977) did not map naturalized vegetation. Serpentine sites on California annual grassland were included under SRM 215 (Valley Grassland).

89. Serpentine Barrens soil- The soil series of the Serpentine Barrens range site just seen is Montara, a Vertisol. Cracking of this azonal soil is due to it's high shrink-swell potential resulting from the high content of montmorillonitic clay (mentioned above) and the shallowness of this upland soil. The physical features of cracking and shallowness together with chemical infertility creates a harsh edaphic habitat for range plants. What is so interesting, however, is that the annual grass species are the same on the Serpentine Barrens as on the non-serpentine site except there are higher proportions of the less palatable species, especially naturalized invaders, on the Barrens. This is because on this range sheep are attracted to and spend more time grazing and, generally, congregating on the Serpentine Barrens sites. The actual reason why sheep preferred grazing and loafing/resting on the barrens was not determined when Rosiere et al. (1986) compared sheep diets from the barrens to diets from non-serpentine sites. Note that even under the relatively heavy degree of use on this site soft chess still set seed for next year.

Foster Pasture, University of California Hopland Field Station, Mendocino County, California. Late May. FRES No. 42 (Annual Grasslands Ecosystem), no Kuchler designation. SRM 215 (Valley Grassland).

90. Comparison of grazed and non-grazed vegetation on a Serpentine Barrens range site- On the left is the grazed Serpentine Barrens range seen in the previous slides. On the right is a 25 year-old exclosure. Naturalized Mediterranean annual grasses dominate grassland communities on both sides of the fence, but there are more individual plants of the native perennial bunchgrass species in the 25-year exclosure (none could be found on the grazed portion). California melic (Melica californica) and purple needlegrass can be seen in the exclosure. Rosiere et al. (1986) concluded that the range vegetation on Serpentine Barrens was not strictly determined by edaphic conditions and features but that it was a combination edaphic-zootic climax. All California annual grassland is disclimax but on serpentine sites the role of grazing appeared even greater. In fact, it seemed (based strictly on personal observation and judgment) that the native bunchgrasses were more common on serpentine than on non-serpentine soils. This is not to say that the native climax grasses would recover and recreate the pre-white man Pacific prairie on serpentine soils if protected from grazing. It does suggest that the native perennial grasses are comparatively better adapted to and more competitive with Mediterranean annual grasses on serpentine sites. Natives like needlegrass, melicgrass, and squirreltail are frequently much more abundant in exclosures on serpentine soils than in exclosures on non-serpentine soils. This is an example of how interactions of management factors (eg. stocking rate, animal distribution) and plant growing conditions influence range vegetation.

Note how perennials extend the green feed season. The annual species have been brown for three weeks and the native bunchgrasses are "still green". On Hopland Field Station the author observed purple needlegrass that retained green shoots through June even on south slopes and through most of July on north and east slopes.

Foster Pasture, University of California Hopland Field Station, Mendocino County, California. FRES No. 42 (Annual Grasslands Ecosystem), SRM 215 (Valley Grassland).

Canebrake

91. Canebrake- The native North American bamboo once formed extensive colonies known to frontiersmen as "canebrakes" along water courses and in flood plains throughout the Southeast from the Pineywoods of east Texas to the Carolinas and as far north as southern Illinois and Ohio. The famous Eighteenth Century naturalist William Bartram described immense canebrakes with phrases like "endless wilderness of cane". In 1819, English botanist and novice explorer Thomas Nuttall described dense canebrakes in bottomlands along the Arkansas, Grand, and Verdigris Rivers, especially at the confluences of these wide streams (Lawson, 2007, ps. 47, 49, 63. 81, 91, 92). Peter Cartwright, legendary Nineteenth Century Methodist circuit rider (and politician who lost to the fledging Arbarham Lincoln) routinely rode for miles of primitive trails (probably of Indian origin) through immense canebrakes that had developed along the floodplain of the Ohio River (Greenbie and Greenbie, 1962, ps. 14-18, 468-469, 478-482).Even as late as the early 1900s Theodore Roosevelt spoke of canebrakes "extending for miles". Canebrakes were the range type that served as the basis of the frontier range cattle industry in areas like the commonwealth of Kentucky (Clark, 1988, p. 158). In his classic, A Winter in the West (Hoffman, 1835, volume II, ps. 164, 210, this latter quoting from the narrative of Colonel Daniel Boone) foot-noted Indians chasing buffalo out of canebrakes in Kentucky. There was apparently much strife between Ohio and Virginia Indians over this hunting ground before appearance of the white man and perhaps giving rise to the common title of a [the] "Dark and Bloody Ground" in reference to wars among the red man even before those between red and white men. Moore (1957) observed that the native bamboo, giant cane or, more commonly, rivercane (Arundinaria gigantea), was so widespread and important as a forage source for livestock and wildlife that to newcomers "Kaintuck", the pronounciation of Dani'l Boone-types, meant "canetuck". When in the severe winter of 1779-1780 giant cane was killed "game perished in sickening numbers" (Moore, 1959, p. 57).

John James Audubon wrote of canebrakes in his Ornitholoical Biography that encompassed the period from 1831 to1839 (Irmscher, 1999, ps. 540-543, 870). Even at that realtively early date in American history canebrakes had already been destroyed over much of their original acreage by clearing for crop fields and livestock overgrazing according to Audubon:

"The Cane, kind reader, formerly grew spontaneously over the greater portions of the State of Kentucky and other Western Districts of our Union, as well as in many farther south. Now, however, cultivation, the introduction of cattle and horses, and other circumstances connectead with the progress of civilization, have altered the face of the country, and reduced the cane within comparatively small limits. It attains a height of from twelve to thirty feet, and a diameter of from one to two, and grows in great patches resembling osier-holts, in which occur plants of all sizes. The plants frequently grow so close together, and in course of time become so tangled, as to present an almost impenetrable thicket. A portion of ground thus covered with canes is called a Cane-brake."

The upper limit of thirty feet may seem exagerated, but Judziewicz et al. (1999, p. 77) gave a height of ten meters (complete with a photograph with shoots approaching that length). Audubon further described canebrakes as "growing beneath the gigantic trees that form our western forests" so that not all canebrakes are strictly grasslands but some are exit as or are components of savannahs or layer(s) in decicuous forests (Irmscher, 1999, p. 540). Audubon (Irmscher, 1999, p. 541) indicted, "The canes grenerally grow on the richest soil, and are particularly plentiful along the margins of the great western rivers". Farmers settled near or adjacent to canebrakes "...because these brakes are plantifully stocked with game of various kinds". He then explained how farmers cleared canebrakes with large knives and burned dried piles of the bamboo shoots which produced loud reports when internodes exploded as moisture was converted to steam.

Schwaab (1973) compiled various quotes for early travels in southern North America including the following in reference to canebrakes in various river bottoms in Arkansas. "In some places are impervious forests of cane, twenty feet high, as thick as they can be stuck" (page 212). "Then you hear a mile off, the continual musketry which cane keeps up in burning, as the air contained in the joints expands and explodes" (page 213). The following short description was in reference to "vast canebrakes" in Kentucky (Schwaab, 1973, p. 267) that were "... tracts of wild cane, very much estemed by the wild and tame cattle, it continuing in verder all through the winter" (p. 55). Probably the most complete collection of descriptions and travelers' accounts of of canebrakes in Kentucky was that of McHargue (1941).

While emphasizing travel in the Ohio Valley, Jakle (1977, p. 115) indicted that by the decade of the 1840s the famed geographer Charles Lyell explained that "[m]any European plants" had become naturalized foreigners that "overrun the country with amazing rapidity". An example of these naturalized European introductions was a bluegrass (Poa pratensis) that it became so well-adapted and so dominant over an area of Kentucky that it lent its name to what became known as the "Bluegrass Region" or the Bluegrass section (Fennemann, 1938, ps. 419, 427-431). For its "payment" this grass was given the specific common name of Kentucky bluegrass. Eventually, Kentucky took for its nickname, "the Bluegrass State". Prior to this agronomic conversion by the whiteman and up until the 1790s the dominant grass in this area had been the native cane, Arundinaria macrosperma [now Arundinaria gigantea] (Jakle, 1977, citing Gilbet Imlay, ps. 115, 190).

To illustrate this change in the major forage grass, your country music-loving author remarked that if Bill Monroe had formed his marvelous musical group in frontier times (in the days of the frontier backwoodsman) instead of in the later settlement and settled era, the stringed instrument troupe been would have more appropriately dubbed, "Bill Monroe and the Canebrake Boys" and the country (hillbilly) musical genre would have become known as Canebrake Music rather than Bluegrass Music. Beef production (largely cow/calf operations) based on grazing tame or agronomic pasture made up primarily of Kentucky bluegrass (and later, of course, Kentucky 31 tall fescue [Festuca arundinacea]) was (is) contemporary stock farming not ranching. Running cattle in the canebrakes was frontier ranching during the open range era.

The most definitive (and certainly the--or one of the--earliest) published treatments of forage resources in eastern North America (at time of its publication, the western frontier of the infant United States of America) was the richly "Germanized" account entitled, A Topographical Description of the Western Territory of North America by Gilbert Imlay (1797). This phenomenal publication of over 600 pages, which included parts of what eventually became Ohio, Kentucky, and Tennessee, was part travelogue and part navigational-agricultural-political-sociological even ecological report. Imlay (1797) described "range", "prairies", "glades", "barrens", "savanna", stock drives, tree-killing/grass-increasing fires, and even sugar maple as cattle browse. Aside from sugar maple the indigenous plant species that received more coverage than any other in A Topographical Description... was "cane" (Imlay, 1797, ps. 29-30, 118, 166, 167, 233, 236, 318, 440, 447). "Cane" so impressed this well-traveled, well-informed, keen observer that he presented morphological details of its shoots and praised it as "perhaps the most nourishing food for cattle upon earth" (Imlay, 1797, ps. 29-30, 318). This assessment of the feeding value of cane was consistent with scientific observations that giant cane provides some of the most palatable forage of any grass species native to North America.

Another rich source attesting to value of canebrakes (probably of both Arundinaria gigantea and A. tecta) on the frontier was that of the self-taught naturalist, Dr. Gideon Lincecum whoses writings were compiled and edited by Lincecum and Phillips (1994). This compilation included various references by Lincecum regarding canebrakes as ideal wildlife habitat and source of livestock forage (Lincecum and Phillips, 1994, ps. 10, 54, 58, 60, 62, 64, 65, 101, 118-119, 170-171). "I consider the canebrake the great sanctum sanctorum; the inner chamber of the great hunting ground"-- Gideon Lincecum apparently in reference to bottomlands of the Tombigbee River in present day Alabama (Lincecum and Phillips, 1994, p. 64).

Canebreaks were vast monotypic stands of giant cane. Unfortunately, canebrakes were quickly destroyed by clearing for cornfields, sugar cane plantations, etc. and by overgrazing of the extremely palatable and nutritious forage by cattle, hogs, and horses (Judziewicz et al., 1999, ps. 77, 197-198). Overburning of this woody grass with its vulnerable perennial shoots was also a factor thought Arundinaria is well-adapted and benefits from periodic episodes of fire (Wright and Bailey, 1982, p. 377-378).

In their encyclopedic Manual of the Grasses of the United States Hitchcock and Chase (1951) distinguished giantcane or rivercane (A. gigantea) from switchcane(A. tecta) on the basis of air canals in rhizomes of the latter. Gould (1975, p. 40) apparaently followed this distinction at species level, but more recent interpretations such as that by Judziewicz et al.(1999, ps. 197-198) recognized two subspecies: A. gigantea ssp. gigantea and A. gigantea ssp. tecta. In volume 24 of the encyclopedia, Flora of North America, Barkworth et al. (2007, ps. 17-20) returned the gigantea and tecta taxa to species level in addition to adding a third Arundinaria species, hill cane (A. appalachiana). The latter was apparently the namesake species of Cane Ridge, the famed location of a multi-denominational camp meeting. Species designations/distinctions in Barkworth et al. (2007, ps. 17-20) were based mostly on such morphological features as number of basal internodes, persistence or deciduousness of culm leaves, and topknots (fan-shaped clusters of leaves at tips of new shoots) as well as the previously used presence or absence of air channels in rhizomes.

These are the only bamboos native to North America and loss of the once vast expanses of this woody grassland from a primeval continent bespeak man's lack of appreciation for grassland vegetation. Millions rally to save a few old-growth specimens of trees but few are so inspired for grasses (even woody ones). Perhaps it is indicative that even the Society for Range Management in it's treatment of North American rangeland types (Shiflet, 1994) totally neglected identification and description of this now-restricted but still extant, and locally important, range cover type.

Canebrakes are so dense and vast as to typically exclude any other plants. However, giant or river cane forms an exclusive understory in both deciduous and coniferous forests. The relict canebrake seen here is along the Brazos River in central Texas where it has a savanna form with sparsely scattered trees of eastern cottonwood, bur oak, pecan, sycamore, red or slippery elm (Ulmus rubra) and box-elder (Acer negundo) throughout the rhizomatous "thickets" of giant cane. Cameron Park, Waco, Texas. October, late estival aspect. Arundinaria gigantea retains its compound leaves throughout the relatively mild winters of its range shedding these in spring with emergence of new leaves. Thus it is an "evergreen" grass. This is probably the main reason why the nutrient-rich leaves are so palatable to large herbivores, and thus why canebrakes were so quickly destroyed by livestock overgrazing. No FRES, Kuchler, or SRM designations. Texas Blackland Prairies- Northern Blackland Prairie Ecoregion, 32a (Griffith et al., 2004).

In recent years--and after decades of neglect--canebrakes were finally recognized ("rediscovered") as comprising one of the most endangered natural ecosystems and native plant communities in North America. Several important projects were begun to save this under-appreciated (previously unappreciated) range type. Restoration of canebrakes finally became a major conservation cause in the United States of America, particularily in the Southeast (Brantley and Platt, 2001). Platt and Brantley (1997) provided an in-depth review of canebrakes in a combination historical and ecological context. One of the major problems in restoration of Arundinaria canebrakes has been--and remains (though to a lesser extent)-- propagation of this predominately asexual reproductive genus of bamboo. Baldwin et al. (2009) described three methods of propagation for A. gigantea, which probably could be extended to the other Arundinaria species. These methods included the more practical and commonly used rhizome planting method, but with more intensive treatment of rhizomes prior to actual planting. The other two methods were elaborate micropropagation procedures and seedling production from elaborate grain germination techniques.

Re-establishment of giant cane "ain't easy". Just another example that it is much better (easier and more efficient in the long run) to take care of something to begin with rather than lose it and then have to resort to Herculean efforts to get back a fraction of what was squandered. In the case of canebrakes it is too late for the proverbial "ounce of prevention", but at least the prodigal sons of the canebrake are coming home.

Introduced (Anthropogenic or Man-made) Canebrakes

Bamboozled Confusion- Several species of exotic bamboos (Baubusae tribe) have been introduced into North America, especially in the southern United States, as ornamental plants. In the most recent monograph of Texas grasses, Shaw (20120 ; ps. 248-250, 801-803) descrbed four introduceed bamboo species (two each in Bambusa and Phyllostachys genera) and cited aources listing 26 Phyllostaachys spceies cultivated in the Lone star State.In the previous monographic treatment of Texas grassses (Shaw, 1974; ps. 40-41).did not list any bamboo species other than the native giant or river cane (Arundinaria gigantea) while mentioning the very similar switch cane (A. tecta).that he regarded as not being in Texas. Likewise, in their coverage of grasses in the Texas Cross Timbrs and prairies Hignight et al, (1988; p.40) gave A. gigantea. as the only bamboo. In the first monograph of Texas grasses (Silveus, 1933; ps. 1-4).listted both Phyllostaachys and Bambusa as well as Arundinaria gigantea inclusing a diagramatic key to separation of the three genera. Borowski and Holmes (1996) made the case that yellow bamboo (P. aurea) has naturalized in Texas (as in Louisiana) often forming dense colonies which may exclude other plants. These workers concluded that yellow bamboo should be recognized as naturalized and, hence, be included as part of Texas vegetation. Diggs et al. (1999, p. 1311) included yellow bamboo in the northcentral Texas flora apparently concluding that it had naturalized or at least persisted and formed large colonies. Given omission of Phyllostachys species from some of the more recent treatments of Texas grasses along with similarity of shoot features to Arundinaria, plus characteristic absence of or infrequent sexual reproduction (Borowski and Holmes, 1996) for positivve identification baed on spikelet morphology, it is not surpising that students of Texas flora and range plant communities (including this author) have been confused by presence of yellow bamboo and frequently regarded it as a native canebrake of A. gigantea. Nonetheless, it can be used a substitute example of a native caenbrake.

92. Canebrake in the Brazos bottoms- As in other southeastern states, Texas was once home to extensive canebrakes, but these are now restricted to relict tracts missed by farmers and protected from overgrazing. The tree trunk here is box-elder, one of the most drought- and heat- tolerant of the maples. The native bamboo in this stand was giant, river, or southern cane (Arundinaria gigantea) Canebrakes are extremely valuable as feed and cover for wildlife, including birds. John James Audubon was renowned for his accuracy and detail is placing his subjects in their habitat. One of his most famous paintings is of the wild turkey (Meleagris gallopavo) in a canebrake. This painting was an art record of the condition described by Cartright (1856) as "that new and beautiful country of 'cane and turkeys'". Judziewicz et al. (1999) discussed A. gigantea at some length and concluded that loss of this vegetation type may have been a factor in extinction of Bachman's warbler (Vermivora bachmanii) and even the Carolina parakeet (Conuropsis carolinensis).

Shown here is part of an established canebrake of yellow bamboo ( Phyllostachys aurea) either planted by human hands, escaped as a waif, or naturalized. This introduced bamboo has the physiogonomy and general structure of a native canebrake of Arundinaria species thus giving viewers an idea of the appearance of North American canebrakes.

Cameron Park, Waco, Texas. October. No FRES, Kuchler, or SRM designations. Texas Blackland Prairies- Northern Blackland Prairie Ecoregion, 32a (Griffith et al., 2004).

93. Canebrake above the reaches of the Bosque- The once vast North American canebrakes of giantcane and switchcane apparently extended some distance above their major habitat of river bottoms. The famous Cane Ridge that figured so prominently in development of backwoods camp meetings and onset of the Second Great Awakening gives some suggestion of this. The above two photographs were of introduced yellow bamboo giving readers an idea of appearance of a former canebrake range community that extended upland for considerable distance from the floodplain of the Brazos River in Texas' Western Cross Timbers. In the concept of Landscape Ecology such former extensive canebrakes along creeks and rivers, perhaps in the Cross Timbers, constituted ribbon-shaped patches imbedded within the matrix of the surrounding Cross Timbers. These waterways and their canebrakes also functioned as corridors connecting grasslands of such prairies as Grand Prairie and woodlands/forests of the Cross Timbers.

Large trees in the background of both slides were post oaks. Smaller trees of hackberry or sugarberry (Celtis laevigata) were in background of the first of these two slides. The tree in left foreground of second slide was pecan. For several years prior to taking of these photogrphs this introduced, and perhaps-naturalized canebrake had been heavily grazed by horses with high-lining" of branches within reach while most main shoots were not consumed. This demonstrated that canebrake range has some resilency.

Erath County, Texas. April. Vernal aspect; shoot emergence/elongation phenological stage. No FRES, Kuchler, or SRM designations. Cross Timbers- Western Cross Timbers Ecoregion, 29c (Griffith et al., 2004).

94. Rapid growth in a favorable spring- Edge of naturalized yellow bamboo ( Phyllostachys aurea) canebrake in mid-spring during development of this year's shoots. Some bamboo shoots had already reached their mature height (first photograph) while limbs were still coming into their final position as short, ephemeral, husklike sheaths were shed to that limbs were released from their appressed position on the culms. This positioning of limbs along sides of culms was shown at outer margin of the canebrake in the second photograph.

Erath County, Texas. May: various phenological stages of shoot emergence and elongation.

95. Shoot emergence of yellow bamboo ( Phyllostachys aurea) - Floor of a naturalized canebrake at onset of annual grass growth cycle. Shoot growth (emergence and elongation of main culm and branches) of giant cane typically occurs only once per year. This is at onset of spring green-up when previous year's leaves are shed and current year's leaves emerge essentially simultaneously. Young shoots at this stage were used as a pot herb by Indians and backwoodsmen. The shoots had to be boiled "for an eternity" (they ain't exactly asparagus).

Erath County, Texas. April: early stage of shoot emergence.

96. The same bamboo shoots as in the previous slide seven days later and now over six feet in height. The outer sheath-like husks that cover the newly emergent shoots are deciduous (ephemerially so) and quickly fall away allowing the branches that were appressed against the culm to drop into place with rapid unfolding of pre-formed leaves.

Erath County, Texas. April: early shoot elongation and sheath-shedding stages of phenology.

97. At 13 days post-emergence the shoots of yellow bamboo in the last two slides have reached their adult (maximum) height of six to 10 feet or more. No more aboveground growth will occur this growing season even though there will be over five months of frost-free temperatures and soil moisture adequate for growth (plus leaves remain green until shed the next spring). Instead all growth occurs underground as rhizomes development and extend for distances of up to 15 feet. Tillers will emerge from these rhizomes the next spring to begin the annual growth cycle again.

"I was raised in a canebrake by an ole mamma lion; cain't no high-toned woman make me walk the line"

(Sixteen Ton by Merle Travis and made famous by Tennessee Ernie Ford).

Prevalence and importance of canebrakes in the Cumberland country as livestock and wildlife feed and habitat and as an impediment to travel was discussed by Arnow (1960, ps. 20, 38, 51, 56, 98, 127, 168, 237, 247, 401, 408) and Arnow (1963, ps. 214-215, 239, 241). Sitton (1995, ps. 6, 7,25, 34, 49, 66, 145, 148, 200, 224, 226, 231) documented canebrakes as forage sources in the open range ranching region of the Big Thicket (deep east Texas) which as discussed under the Forest and Woodland Slides (oak-hickory range type) was the last actual open range in the United States. Blevins (2002, ps. 18, 27, 35) explained that in the Arkansas Ozarks switch canebrakes either became some of this states richest farmland or were used as open range by cattlemen during roughly the last two-thirds of the Nineteenth Century. Beilmannn and Brenner (1951, p. 276) reported from a historical review of travlers' accounts, explorers' diaries, and especially surveyors' notes that the "impenetrable" canebrakes were a "popular retreat of game of all sorts", especially "one of the favored haunts of the black bear". One of the most interesting (and least remembered) "habitat" uses of canebrakes was as a hideaway for runaway slaves, renegade Indians, missionaries, and southern "white trash". Accounts by Peter Cartwright, one of the most influencial and colorful Methodist circuit riders in America, described villages made up of mixed-blood misfits and social outcasts hidden in the immense canebrake that streatched for miles along the floodplain of the Ohio River (Greenbie and Greenbie, 1962, ps. 14-18, 468-469, 478-482).

"Raised in a canebrake" may have been a frontier expression (much like "born in a barn" as for example, "born-in-a barn Arkansawyer"). It might also have been a backwoods claim of honorable and humble birth (the two being one and the same among backwoodsmen) similar to being born in a log cabin. Apparently, Brother Cartright claimed that he had been born in a canebrake, the same canebrake where his mother had hidden in an Indian attack (Bray, 2005, ps. 1, 277). Rev. Cartwright was, afterall, an astutre politician and one who once ran against a candidate who had been born in a log cabin, Abraham Lincoln. (Incidentially, from his own autobiography [Cartright, 1856] and the historically valid account of his life (Bray, 2005), Rev. Cartrighrt also knew quite a bit about "high-toned women".)

98. Advance of a canebrake- Invasion of "new land" by esotic (and now apparantly naturalized) yellow bamboo. Shoots (tillers) from remarkably long rhizomes produced over course of the previous growing season formed the vanguard as a canebrake advanced into ground not previously colonized (at least not in recent years) by this native species of bamboo. Newly emerged shoots were in various stages of development. Most shoots had just shed the "husks" from their culms thereby permitting the until-then appressed limbs of the shoots to be released and attain their mature positions along the culms. These shoots ranged in height from less than one foot to over six feet in height.

Erath County, Texas. May: various stages of shoot emergence and early development.

99. Shortleaf pine-oak forest with a canebrake understorey, a pine-oak-giant cane savanna- This was a most unique composite vegetation. It was an oak-pine forest with tree and shrub layers that were diverse in their own rights to which was "added" a canebrake of giant, river, or southern cane, the native bamboo (Arundinaria gigantea subsp. gigantea). There was no layer of vegetation (at least of macroscopic vascular plants) beneath the canebrake which is the typical condition for canebrakes of this species. Transeau et al. (1940, p. 762) specified that canebrakes were "often invaded" by evergreen oaks. As described in the next paragraph evergreen species in this community were pines and the oak species were not "live oaks". This community was similar, however, to that described in a standard Botany text widely used decades ago (ie. old textbook knowledge). Prior to settlement by European man, giant or river cane formed vast "brakes" or colonies (as in endless miles of bamboo) along water courses ranging in size from major rivers to creeks throughout southeastern North America. These seemingly infinite canebrakes occurred as far north as the Ohio River and westward to central Texas. Giant cane also grew (and still grows) as isolated plants of one or a few shoots up to small colonies of several hundred square feet as part of the understorey vegetation in oak-hickory and pine-hardwood forests. An exclusive (a homogenous or "pure") stand of uninterrupted canebrake understorey beneath an adult tree canopy was-- in this vegetation hunter's experience-- quite rare. The species composition, structure, and physiogonomy of this forest-woody grass vegetation was likely that of the climax. This was relict vegetation. Even though it was not virgin forest this was interpreted as a climax community tht lacked the component of very old trees of the species present. It was one of the most unique assemblages of native plants encountered by your photographer-author.

Shortleaf pine was the dominant tree, but there were a few individuals of loblolly pine (or perhaps of the widespread natural hybrid of these two species). Other trees included white oak, southern red oak, water oak, and sweet gum in about that order. There was a shrub layer composed of small individuals of persimmon and sassafras as well as smooth sumac. These shrubs and/or short trees grew in the canebrake, but were shorter than the taller shoots of the bamboo except at the edge of the canebrake where there was light of greater intensity and frequency. Pensacola bahiagrass (Paspalum notatum var. saurae) and hurrahgrass (P. pubescens= P. muhlenbergii= P. setaceum var. muhlenbergii) also grew on these extreme edges where they got adequate light for survival.

The development of spreading, horizonal branches on all trees, including the dominant pines that are usually strongly apically dominant, indicated that these trees had developed on a savanna which was exactly what this community appeared to be: an oak-pine-giant cane savanna. The soil was moist enough to support water oak and the scarce loblolly pine (or a hybrid thereof), but soil water conditions were not periodically ponded let alone swampish.

The grassland expression of Arundinaria gigantea canebrake (exclusive, homogenous stands or colonies of giant cane with widely scattered or infrequent trees) was covered in the Miscellaneous portion of the Grasslands section.

Miller County, Arkansas. July. West Gulf Coastal Plain portion of the Coastal Plain physiographic province. Pineywoods vegetation, part of the Southern Pine Region. According to U.S. Forest Service Ecological Divisions (Forest Service, Final Environmental Impact Statement, 1999) this was in the Southeastern Mixed Forest Province (231), Mid Coastal Plains, Westen Section (231E), South Central Arkansas Subsection (231Ea). FRES No. 14 (Oak-Pine Forest Ecosystem). K-102 (Southern Mixed Forest). SAF 76 (Shortleaf Pine-Oak, variant of). No Kuchler or SRM units for canebrake. South Central Plains- Tertiary Uplands Ecoregion, 35a (Woods et al., 2004).

100. Shortleaf pine-mixed oak-giant cane savanna or a shortleaf pine-mixed oak forest with a giant or river cane understorey- A more interior view of the composite pineywoods vegetation displayed in the preceding slide. Interpretation of this vegetation was subject to many subjective factors including disciplinary background or professional allegiance. It could be viewed as a mixed oak-southern pine forest with an understorey comprised exclusively of a "brake" of giant cane, the one species of native North American bamboo. Alternatively this could be viewed as a combination of diverse pineywoods forest and giant cane or bamboo grassland. Beyond any debate these two "elements" (components) were present. Also beyond debate is the ecological fact that both of these vegetational elements do occur (they formerly occurred on massive scales in the virgin vegetation) in their own forms separate and distinct from each other as potential natural vegetation. It was a different form of canebrake; it was a unique developmental expression of oak-pine forest growing as it were "out of" or in the midst of a canebrake.

The characteristic leaves of southern red oak appeared very distinctive along both the left and right borders of this photograph. Smooth sumac and shrublike or scrubby trees of persimmon and sassafras were overwhelmed in a "sea" of native bamboo.

Miller County, Arkansas. July. West Gulf Coastal Plain unit of Coastal Plain physiographic province. Forest Service ecological divisions given in preceding caption. FRES No. 14 (Oak Pine Forest Ecosystem). K-102 (Southern Mixed Forest). SAF 76 (Shortleaf Pine-Oak, variant of). No Kuchler or SRM unit for canebrake. South Central Plains- Tertiary Uplands Ecoregion, 35a (Woods et al., 2004).

The following section on canebrakes in the western section of the Ozark Plateau, which is the Springfield Plateau, was based on field observations and photographs (35 mm slides) taken in McDonald County, Missouri on the first day of winter. The author was "carried" by his fellow faculty member and "soul mate", Dr. William R. (Bill) Stevens, to two populations of giant cane (Arundinaria gigantea= A. giganteasubsp. gigantea) in two locations (one on Elk River, one on Little Sugar Creek). The author was about half-way trespassing when he took the following slides (he had belated permission on one canebrake location and indirect permission to enter the other canebrake), but once a few local folks came poking around Bill got edgy about the author's photographic safari (even though Bill had volunteered to provide the transport in ole Precious, his half-ton Ford pickup).

At the second stop a nosey neighbor came up to Bill (who had remained in Precious while his photographic friend was "shooting canebrakes") asking if the "trespasser" had permission from "Pat" to go on "Pat's land". Bill told Nosey that he was not for sure. (Your photographer had just gotten permission from one of "Pat's" extended family members at the first "photographic opportunity".) Upon getting an indecisive answer from Bill the nosey McDonald Countian, who obviously saw himself as Deputy Barney Fife (and in reality was a counterfeit or mere would-be hillbilly), tore off to warn "Pat" that somebody was snooping around on his land. Your author/photographer got back to Precious in due time (he opportunistically did not hear Precious' loud horn honking). Thereupon, Bill made due haste to leave McDonald County, Missouri and its remnant canebrakes behind to return to "civilization" in Newton County, Missouri (the northern adjoining county) where presumably landowners were a little less apt to shoot first and ask questions later.

Or course, Bill's photographic canebrake tourist was wanting to meet "Pat" because the apparent landowner might know of more canebrakes. Your author, who fashioned himself a bona-fide hillbilly, was confident that "Pat" would be proud that his canebrake was making a contribution to science and education. Most landowners are decent when it comes to such-- even if they "don't known what it's all about it". Bill indicated that there could be "shootin' exceptions".

The next evening, after an afternoon of getting Christmas pecans and touring the northeastern Oklahoma countryside (tame pastures in particular) in Precious, the sixty seven-year-old Bill Stevens died while talking on the telephone to this author. Cause of Bill's death remained unknown, but appeared to be cardiac arrest, stroke, or some form of respiratory failure. If this had happened in Precious earlier that day touring Green Country Oklahoma or the afternoon before while outrunning "Pat"...

The following section was dedicated to Dr. Bill Stevens, Professor of Business Administration, Missouri Southern State College (later, University). This was the only part of Range Types of North America�dedicated to any individual person. To have done less would have been heartless and dishonorable. So here's to Bill-- for some last heartbeats and lasting happiness in Ozark canebrakes (and more fond memories from a lot of other times and places). So long, loyal friend.

101. What the riverside used to be- A relict tract of canebrake--a population of giant or river cane, the native bamboo of the Ozark Plateau,-- growing on the immediate bank of Elk River in southwest Missouri on the first day of winter. This remnant of climax natural vegetation extended from the riparian zone of Elk River up the slope of an adjoining hillside. There were trees growing in/on/through the canebrake such that this natural vegetation was a savanna form (a savanna rather than a grassland per se). Trees in these scenes of a savanna form of canebrake were mostly huge (compartively humongous; nearly two foot diameter breast height) specimens of honey locust (Gleditsia triacanthos) with some sapling to pole-sized white oak (Quercus alba).

This canebrake provided a textbook example of a reference area of a vegetational relict or relict vvegetation: a remaining tract of the original native vegetation (= the natural plant community that existed just before or prior to impact of technologically modern man). Such reference relict tracts are typically relatively small in size compared to total extent of the original plant community, but they serve as a representative sample or example-- a reference stand--of the virgin vegetation, the plant community before it was modified by Homo sapiens.Relict vegetation in/of an ecological reference area is usually assumed to be the climax (potential natural) vegetation�except where the habitat (= range or forest site) of the plant community and thus, the present plant community, is still undergoing plant succession and, hence, cannot have development of a climax community. Sere�(Clements, 1916) is the pathway or pattern of vegetation development for/on a given�site.

By common usage in Vegetation Science, relict and reference are used synonymously when applied to tracts, areas, and sites (seres) even though there are slight (and, perhaps, strained) differences among them. (Of course, highly degraded tracts, areas, sites, etc. with abused and deteriorated vegetation can also serve as�reference �tracts though obviously they are at the opposite end of plant succession on a given site). Other terms for reference areas include natural areas, ecological (or ecosystem) reserves. or climax tracts among others (Laycock, 1975). By the way, the definitive "reference" for range reference tracts is Laycock (1975).

Tracts of�relict�plant�communities�(especially if they are of climax vegetation) serve as "bench marks", "base lines", "high water marks", etc. of what the plant life of a given�site, as a plant community, should be on that site if this natural or native habitat was not substantively modified by man (ie. if it still had itsnatural sit potential).

Relict or reference vegetation serves as ecological (specifically, successional) evidence of change (dynamics) or, alternatively, of stability in the natural plant community.Hence, vegetational relicts function as the equivalent of "botanical forensics". Florensic(s) has been defined variously, but florensic essentially means or refers to the application of scientific concepts and procedures (ie. knowledge) to determine facts (= truth) by standards established by--and, thus, as admissible evidence in, certain spheres of inquiry especially in courts of law (as in criminal cases before the bench). Thanks to television dramas, in common usage or popular culture florensic evidence implies analysis of blood, tissues, gun powder, hand-writing, and murder weapons. There is the equivalent for "ecologicl crimes" to vegetation (as in the speciality of Range Analysis for example) with florensic methods including plant identification, soil and plant chemical determination, even "motive" (= management objectives) and vegetational "wounds" or "tissue damage" by cutting edge weapons like plows, axes, teeth, hooves, and fire.

Relict vegetation or reference areas serve as the basis or the standard by which to measure the degree or extent of damage induced by the "crime" (tillage, overgrazing, improper application [or non-application] of fire, "cut-and-run" logging) that induced, contributed to, or caused retrogression (retrograde movement down a sere) of natural vegetation. Such was the case with the relict canebrake featured in this section. The two canebrakes shown and described in this portion were ecological (successional)--and incontrovertable--evidence that the native or natural, the pre-whiteman, vegetation�along Elk River and Little Sugar Creek of the Springfield Plateau in southwest Missouri was a canebrake of giant or river cane. The savanna form of canebrake seen in this section was the riparian vegetation (and the adjoining hillslope vegetation) of Elk River and Little Sugar Creek before the whiteman destroyed it.

Sadly, today much, if not most, of the banks of Elk River are piles of river gravel largely devoid of much of any vegetation. A very sick and sickening scene; a stark commentary of the whiteman's lack of stewardship of one of the most precious natural habitats stolen from the redman.

How this small relict of climax riverbottom vegetation had persisted was probably lost to local oral tradition and now known but to God. While your author was taking these (and other slides in this set) a ravishingly beautiful, young woman in heavy green Christmas socks (a winter adaptation of bare feet) came out to the apparently trespassing photographer to inquire about his activity. Your author introduced himself as being a native of the adjoining county and now a professor in Range Management who was collecting photographic material for his on-line, web publication. The ravishing hillbilly, who lived in a semi-secured trailer-house thirty yards away (and about the same distance from the bank of Elk River) became all the more a lady and told your author what she knew about the canebrake including the location a a larger one along Little Sugar Creek. That property was owned by the same family as the land the interlopering photographer was now treading (lightly) upon. After a while the two Ozarkers parted company after exchanging "Merry Christmas" and your excited rangeman set off in search of the other canebrake. (This author wished afterward that he had asked the Christmas-socked lady to stand by some of the bamboo. Just for scale you understand. So much for a canebrake in hindsight. Well, maybe a for-scale shot or two next summer-- and without socks or long pants.) To be continued...

Elk River, McDonald County, Missouri. Late December, hibernal aspect. FRES No. 14 (Oak-Pine Forest Ecosystem). K-102 (Southern Mixed Forest).No SAF for this form or variant of an Ozark Plateau bottomland forest with an understorey of giant cane. No Kuchler or SRM units for canebrake. Ozark Highlands- Elk River Hills Ecoregion 39b (Chapman et al., 2002).

102. Just beyond the river bank- A canebrake of giant or river cane just back from the immediate bank of Elk River in the Springfield Plateau of southwest Missouri. The large trees (up to two foot diameter breast height) were honey locust, an Intolerant species that--like sycamore and cottonwood--thrives on the water-disturbed river bank as a pioneer which then persist to old age and great size in the climax vegetation.

This climax vegetation could arguably be regarded as a bottomland forest with an understorey of giant cane that formed a canebrake as a lower vegetational layer. Alternatively, this range plant community could be interpreted as a canebrake with enough cover and density of trees as to be a savanna form of canebrake. The author opted for this latter vegetational interpretation given that plant cover, density, standing crop, and biomass production was far greater for grass than for trees so that this climax range vegetation was grassland and clearly not forest or, even, woodland (not by any proverbial stretch of the vegetational imagination).

Note that in this regard the riparian vegetation was overwhelmingly grass not tree or shrub. This was a climax grassland, specifically a canebrake with sufficient tree cover and density as to result in a physiolonogy of a giant cane-tree savanna.

* That canebrakes covered immense stretches along numerous rivers and their tributaries in the western Ozarks prior to modifications by the whiteman is a fact documented by early travelers and, especially, explorers. The most prominent of these explorers was Henry Rowe Schoolcraft who (while accompanied by Levi Pettibone) explored the western Ozark area for four months from late 1818 through early 1819. These inexperienced yet keen observers wandered throughout the western Ozark Highlands (=Springfield Plateau) of southern Missouri and northern Arkansas keeping a detailed journal of their nearly misadventure. From this minimal (adjectives such as poor-boy, ill-equipped, greenhorn, tenderfootwould be understatements) expedition came the classic, the exploratory masterpiece, Journal of a Tour into the Interior of Missouri and Arkansaw (Schoolcraft, 1821). This first written (at least, first published) account of exploration of the Ozark Region was republished, along with a synopsis of the "ordeal", by Rafferty (1996) under the primary title of Rude Pursuits and Rugged Peaks. All true-blue Ozarkers and would-be hillbillies must read this account (under one title or the other). Schoolcraft (1821) left many entries in his journal of extensive canebrakes along Springfield Plateau streams. The extensiveness and nearly impenetrable botanical barrier of such canebrakes obviously impressed him. One note of caution for readers of the Rude Pursuits and Rugged Peaks edition is that in the synopsis, which was written by Rafferty (1996) and who was obviously not a biologist, woodreed (Cina arundinacea)--a native, herbaceous, climax bunchgrass of bottomland forests--was frequently confused with giant or river cane (Arundinaria giganta).

More recently--and to this author's knowledge the best summary account of canebrake range in the Ozark Uplift--Blevins (2018, p. 146) provided the following historical perspecctive on use of Arundinaria gigantea for forage by frontier backwoodsmen: "...In wintertime, herders sought out extensive canebrakes to avoid the expense and hassle of putting up hay and feeding it. Pioneer settlers recalled stands of Arundinaria gigantea reaching truly gigantic heights of twenty feet on river banks and in the alluvial bottoms of streams. Cattle and wildlife feasted on switch cane's evergreen leaves. So crucial was the canebrake to successful cattle raising that herders developed unwitten codes to govern its use in the say way that codes mediated homesteading, bee tree discoveries and other informal functions of pioneerlife. 'Early settlers had a custom which they strictly observed', remembered one man of the upper White River valley in the 1830s. 'Each settler would select a bottom on a creek or the river, that had a canebrake on it, to winter his cattle and horses, and no one else would dare to turn stock on it'".

Outer bank and first terrace of Elk River, McDonald County, Missouri. Late December, hibernal aspect. FRES No. 14 (Oak-Pine Forest Ecosystem). K-102 (Southern Mixed Forest). No SAF for this form or variant of an Ozark Plateau bottomland forest with an understorey of giant cane. No Kuchler or SRM units for canebrake. Ozark Highlands- Elk River Hills Ecoregion 39b (Chapman et al., 2002).

103. Just above the floodplain- An upland stand, a population on a low hillside, of giant or river cane that formed a dense canebrake beneath scattered second-growth white oak (ie. a white oak savannah form of canebrake) in the Elk River Hills of the Springfield Plateau in southwest Missouri. This local relict range plant community (not more than two acres in extent "at the outside") was an example of both 1) a canebrake, a savanna form (in this instance) of climax woody grassland that in pre-whiteman times stretched for miles along numerous Ozark Plateau rivers and 2) a reference natural area or relict native vegetation. Shoots of giant cane varied in height from two and half feet to over ten feet in height.

The author first saw this relict stand about eight to ten years prior to the taking of these photographs. Over that time period the following changes occurred: 1) area in canebrake increased (to at least one and a half times), 2) density of cane shoots increased, and 3) height and diameter of cane shoots increased substantially. White oaks had also conspicuously increased in size.

Stands of giant cane once occupied thousands of acres along watercourses throughout the Ozark Region. In his classic Flora of Missouri, Steyermark (1963, p. 81) stated that vast acreages of giant cane along White River and its tributaries had been destroyed by man-made multi-purpose dams like Table Rock and Bull Shoals. This phenomenon of vast canebrakes along creeks and rivers also existed far to the north (and south) of the Ozark Plateau including major rivers and their tributatories such as the Mississippi, Ohio, and Tennessee Rivers. Immense canebrakes once grew along portions of Chesapeake Bay, including its Great Dismal Swamp.

Published distributions of the species range of�Arundinaria gigantea�varied depending on whether this taxon was interpreted as�A. gigantea�(in which case�A. tecta�was a separate species) as treated by some taxonomists and agrologists (Fernald (1950, p. 96; Hitchcock and Chase, 1951, ps. 27-29;�Barkworth et al., 2007, ps. 17-20), or as�A. gigantea ssp.�gigantea�(and with�A. igantea�ssp.�tecta�included in the species resulting in a larger and less restrictive biological range) as viewed by other agrostologists (McClure and Soderstrom, 1973, ps. 26-27; �Kucera, 1998, ps. 25-26). Generally, however,�A. gigantea (=�A. gigantea�as distinct from�A. tecta) and/or A. gigantea� ssp.� gigantea�(as distinct from�A. gigantea�ssp.�tecta) grows over a large region of North America that is bounded as far north as northern West Virginia and southern Virginia and as far west as eastern Oklahoma and Texas. Finally�McClure and Soderstrom, (1973, ps. 25-26) explained that there were hybrid populations of�A. gigantea�ssp.�gigantea,�A. gigantea�ssp.�tecta, and �A. gigantea� ssp.� macrosperma �(this latter subspeies was a new status proposed in the publication of�McClure and Soderstrom, [1973, ps. 25-26]).

While the perimeters of the species range of giant or river cane may be about as extensive as in "Indian times", the once-vast cedarbrakes of this magnificent species were all but eliminated through tillage (conversion to farmland), improperly timed fire, and overgrazing with livestock. Various valiant efforts have and continue to be exerted to restore canebrakes, as for example (Zaczek et al., 2004), so that at least some of the former grandure of the native woody grass and its unique community can be experienced by future generations.

Base of slope immediately above Elk River, Elk River, McDonald County, Missouri. Late December, hibernal aspect. FRES No. 14 (Oak-Pine Forest Ecosystem). K-102 (Southern Mixed Forest). No SAF for this form or variant of an Ozark Plateau bottomland forest with an understorey of giant cane. No Kuchler or SRM units for canebrake. Ozark Highlands- Elk River Hills Ecoregion 39b (Chapman et al., 2002).

104. Deeper on the hillside- Interior of a canebrake of giant or river cane, the giant bamboo native to the Ozark Plateau, growing on a hillside (base of a hill slope) just above the floodplain of Elk River in southwestern Missouri. Leaf litter on the ground surface was comprised of shed, modified leaf sheath coverings, ephemeral bracts that serve as a temporary (just one to three days or so) outer sheath on emerging shoots. The author could not find any other plant species present in the interior of this canebrake.

Base of an east slope immediately above Elk River, McDonald County, Missouri. Late December, hibernal aspect. FRES No. 14 (Oak-Pine Forest Ecosystem). K-102 (Southern Mixed Forest). No SAF for this form or variant of an Ozark Plateau bottomland forest with an understorey of giant cane. No Kuchler or SRM units for canebrake. Ozark Highlands- Elk River Hills Ecoregion 39b (Chapman et al., 2002).

105. A canebrake in late winter afternoon- Exterior of a local canebrake of giant or river cane that developed along a stream (a typical western Ozark Plateau creek) not far from its confluence with Elk River in southwestern Missouri as seen of the first day of winter. This was a savanna form of canebrake with trees mostly of hackberry or western hackberry (Celtis occidentalis) and, secondly, of white ash (Fraxinus americana). On the perimeter of this canebrake there were some plants (or, more precisely, dead weathered shoots) of panicled or panicleleaf tick-trefoil (Desmodium paniculatum). Essentially no other plant species were present in this understorey consociation of river or giant cane.

Scene in late afternoon (roughly two hours before sundown) on the first full day of winter right after a "blue norther", an Arctic high-atmospheric pressure air mass, brought in dry, crisp air. The result was a cobalt-blue sky with just a few remaining cumulus clouds. Perfect contrast for the still autumn-green coloration of giant canes's distinctive�evergreen�leaves. (Some cane leaves had senesced, meaning completed their life cycle and died, which resulted in the "spots" or "small patches" of tan-coloration in these images.)

This relict reference site was a recovering remnant of giant cane that grew on the bank and immediate floodplain of a typical western Ozark Highlands stream which was the edge of a tall fescue (Festuca arundinacea) hay field. Absence of dung, spot grazing, salt feeding areas, cattle trails, etc. showed that no livestock had grazed the stand of tall fescue, and--judging by vigor and size of giant cane shoots--it could be safely inferred that such had been the case for a number of years. Livestock would have grazed-out the giant or river cane long before turning to the unpalatable, toxic (endophyte-containing) Kentucky 31 tall fescue.

Outer bank and first terrace of Little Sugar Creek, McDonald County, Missouri. Late December, hibernal aspect. FRES No. 14 (Oak-Pine Forest Ecosystem). K-102 (Southern Mixed Forest). No SAF for this form or variant of an Ozark Plateau bottomland forest with an understorey of giant cane. No Kuchler or SRM units for canebrake. Ozark Highlands- Elk River Hills Ecoregion 39b (Chapman et al., 2002).

106. A winter afternoon in an Ozark canebrake- Two views of the exterior of a local canebrake of giant or river cane that developed on the far bank of a typical stream in the Springfield Plateau in southwest Missouri. This was a consociation of giant cane that developed as a savanna form of canebrake; that is, as the understorey of bottomland vegetation in which young second-growth trees of hackberry (the dominant tree species) and white ash (the associate tree species) completed the savanna (gave the woody grass grassland a savannah physiogonomy). Some observers might interprete this climax range plant community as a woodland given that tree crowns did not interlock to form a forest yet were closer together than in more typical forms of savannah. The observer who took these slides and traversed this climax plant community interpreted this bottomland vegetation as a grassland not a woodland or forest given that plant biomass and, apparently, gross primary production was overwhelmingly that of giant cane and not of tree source. If this range vegetation was grassland (as seemed indisputable) then the range plant community had to be savanna and could be neither woodland or forest, two forms of vegetation which by definition require(s) a predominance of plant cover, density, productivity, standing crop, etc. to be that of trees.

These views were in late afternoon (roughly two hours before sunset) on the first full day of winter under crisp, dry air and an azure-blue sky imported by a "norther", an Arctic air mass, with a few lingering cumulous clouds. Such was a perfect atmospheric "backdrop" for the late autumn-green leaves of giant cane which, in these slides, appeared in their natural coloration. Some cane leaves had died at end of their life cycle. This phenomenon was responsible for small "patches" (micro-areas) of tan- or light-yellow color in this canebrake. Details of giant or river cane shoots were given in a later portion of this treatment.

107. Green in an Ozark winter- Near exterior of a canebrake of giant or river cane on the bank and immediate floodplain of a creek in the Springfield Plateau of the Ozark Highlands or Ozark Plateau. These two slides presented the "outer edge" or perimeter of a remnant stand of what can conclusively, unequivocally be mentally reconstructed as a former extensive canebrake in the Elk River Valley of southwest Missouri. This series of slides presented a photographic examination of relict vegetation that served as a range reference area of what, in the virgin Ozark Plateau, was an edaphic/topographic climax plant community that extended from the riparian zone of streams upslope to nearly the top of adjoining hills, the roots of former mountainous mountains, in what is called the Elk River Hills.

This remnant canebrake was a long, relatively narrow band of climax vegetation that had survived (it had undoubtedly recovered from a past history of abuse including logging, overgrazing, and, probably, repeated spring burning) on the bank of Little Sugar Creek which was a tributary of Elk River (the confluence of these streams was less than a half mile distant). It can be concluded that in pre-whiteman time (centuries, millenia, etc.) there was a continuous canebrake along the corridor of Elk River and its several tributaries (ie. much of the floodplain of Elk River Valley was a canebrake).

Our photographic excursion--our Nikon laboratory field trip--has thus far included 1) a relict tract of canebrake that was the riparian vegetation of Elk River and, thence, up an adjacent hill as a river cane-white oak savanna form of canebrake and 2) a second relict or remnant tract of canebrake that was a giant cane-hackberry-white ash savanna. The second relict canebrake was shown in these two vertically oriented slides. It was explained above that this climax range vegetation was primarily a stream-bottom grassland and not a forest or woodland because the overwhelming predominance of plant biomass, cover, density, and standing crop productivity was of the native bamboo and not that of indigenous trees or shrubs.

As was the situation for all photographs in this series, these two slides showed this climax range plant community under the bright afternoon sky (about two hours before sunset) following a "blue norther" (an Arctic air mass that developed into a large, atmospheric cell of high pressure) on the first full day of winter. Color of cane shoots seen here was "pretty much" their natural coloration.

Our next photographic stop is entry into the interior of this relict canebrake.

108. Deep inside- Interior of an Ozark canebrake of giant or river cane in early winter on the outer bank of a stream in the Springfield Plateau of the Ozark Highlands. This population of Arundinaria gigantea�shoots was dense (hence, shaded enough) that no other plant species were growing therein. Dead leaves on the soil surface were leaves of�A. gigantea�from the previous spring as well as from a few other previous years. Most of the remaining (incompletely decomposed and still "raw") leaves were the short sheaths (small, modified bracts) that had covered shoots, especially the branches---lateral secondary shoots--that had been appressed (pressed flat against) the main shoots of giant cane. (Morphology of cane shoots was shown and described below.)

There were no other plant species to be found in the deep interior of this canebrake. Plants (weathered, dead shoots) of dormant panicled or panicleleaf tick-trefoil were prewent at the outer edge of this canebrake, but there was nothing other than cane shoots inside the canebrake.

Technical note: less than perfect clarity and limited depth of field in these two slides were a consequence of having to take these shots at 1/15 second in late afternoon light inside this "bamboo drapery".

109. Outer edge of an Ozark canebrake- Exterior of a population of giant or river cane that developed on the outer bank of Little Sugar Creek, a tributary of Elk River, in extreme southwest Missouri. This relict tract of an edaphic/topographic climax grassland (there was enough cover of second-growth hackberry and white ash to form a savanna physiogonomy) was a botanical "Last of the Mohigans" (James Fenimore Cooper) canebrake. Before arrival of pale-skinned human Ozarkers the virgin vegetation of this canebrake probably extended from the riparian zone of streams outwards for considerable distance (perhaps nearly to the top of adjoining low hills) in the Elk River Valley.

Various written accounts of the Ozark Plateau just before permanent entry and establishment of whiteman--such as Journal of a Tour into the Interior of Missouri and Arkansaw (Schoolcraft, 1821)--left no doubt as to the widespread and prominent occurrence of canebrakes along streams in this physiographic province. Later treatments such as the Flora of Missouri (Steyermark, 1963, p. 81) also reported large acreages of canebrakes formed by populations of river cane in the Ozark Highlands. Currently only a fraction of one percent remains of this once dense, luxurant, woody grassland.

These were back-glance views captured after leaving the interior of the canebrake (two immediately preceding slides) on the first full day of winter under a cobalt-blue sky courtesy of a "blue norther" Arctic air mass.

Shoot details- The following (next) series of slides presented shoots of giant or river cane that grew in the two relict tracts of canebrake along Elk River and Little Sugar Creek in the Elk River Hills/Elk River Valley of southwest Missouri (McDonald County). These slides, which were taken on the first full day of winter under a bright blue sky, presented cold weather-hardened shoots with their typically evergreen compound leaves.

110. An Ozark bamboo curtain-�Tillers�of giant or river cane showing the general habit (morphological features) of these vertical aboveground units. All of the�Arundinaria�species grow vast underground networks of rhizomes, subterranean horizontal shoots, as well as their fibrous root systems. Students should remember that the two basic units of the grassshoot�are the 1) culm�or the specialized jointed grass stem with its node-internode unit (the phytomer) and 2) leaves (with their various units including the main parts of blade nd sheath).

Each of these tillers is a module, clone, or ramet�of the original genetic plant (the genotype; genotypically unique individual) that arose from sexually produced seed (recombination of deoxyribonucleic acid from sperm and egg cells). The genetic individual is the genet�with each shoot,�either tiller or rhizome, being a ramet, module, or clone of the genet. This pattern of repeating ramet�production is asexual reproduction which, in the case of cane, results in a canebrake. In a canebrake which tiller belongs to which genet (and what/where that original shoot, the primary or originating-from-embryo shoot is) remains something that only God could figure out. All subsequent ramets of the priary shoot constitute secondary shoots. Likewise, only DNA testing could determine how many genets, whether only one or several, make up a canebrake, all of which have by now been greatly reduced, when not completely eliminated, by human action such as clearing for tillage, overgrazing, inundated by creation of man-made lakes, etc..

In these slides, cane shoots were more or less their natural color when photographed on the first full day of winter under a beautiful azure sky produced by a "blue norther", a high-pressure Arctic air mass. It was explained above that senescence, death at end of the life cycle, of some cane leaves resulted in local spotsof tan or light-yellow coloration on some tillers.

At the size of these slides as seen in this publication, details of tillers were not clear, hence tillers were shown at closer camera distance in the next slide/caption unit. Next slide please...

Outer bank and first terrace of Little Sugar Creek, McDonald County, Missouri. Late December, post-peak standing crop, winter semi-dormancy stage.

111. Cane shoots- Closer-in views of tillers, upright or vertical grass shoots, of giant or river cane showing culms and leaves of these vertical shoots�growing in a relict parcel of canebrake. Visible in these two slides were branches off of the culm, the jointed grass stem, with as few as two to as many as seven leaves on both primary branches and/or their secondary branches. Details of the arrangement of leaves of giant cane were presented farther below. Each cane tiller is a remet, a module or clone, of the original genotypic individual, the genet.

The canebrake that was comprised of these tillers extended from the riparian zone of Elk River outward beyond the river's floodplain and extending almost to the low summit of adjoining hills, some of the Elk River Hills, in the western Ozark Plateau, Springfield Plateau portion or section of the Ozark Highlands. These tillers, vertical or upright shoots, and their evergreen leaves were photographed on the first full day of winter. The green color of leaves and the purple color of culms was essentially the natural coloration of these organs.

Some of the better-known and more thorough descriptions of shoot features of�Arundinaria gigantea�were those of Hitchcock and Chase (1951, ps. 27-29) and Barkworth et al. (2007, ps. 17-19), but the "absolute best", (though much less well-known, treatment of�Arundinaria�species in this regard remains that in the classic treatise of bamboo species native to the Americas by McClure and Soderstrom (1973, ps. 23-46).

Outer bank and first terrace of Elk River, McDonald County, Missouri. Late December; post-peak standing crop, winter semi-dormancy stage.

112. Upper parts- Upper portions of shoots of giant or river cane in a remnant or relict canebrake on the far bank (first terrace) of Elk River in the Springfield Plateau of southwest Missouri. Arundinaria�species are the only woody grasses native to North America. They are also the only North American grass to have petioled or petiolate leaves. The rudimentary petiole of �Arundinaria �species is technically a pseudopetiole� (Barkworth et al. [2007, ps. 17-19]). Leaves of�Arundinaria�species are usually evergreen (barring extremely frigid temperatures) and these leaves (as well as very small, just-emerging shoots) are extremely palatable to grazing wildlife and livestock. Typically there are (two) three to, rarely, seven leaves--with four or five being the most common number--growing on each primary�or, more commonly, on secondary branches..

Primary branches come off of (originate from) the culm. The first branch unit (the primary branch frequently branches again to form secondary branches with each secondary branch bearing two up to, perhaps, as many as seven leaves. In this way, leaves of giant cane appear as if growing in groups or, a shoot apex tips (apices), as clusters of leaves which, as described below, are known as tiller topknots. This pattern was shown in figures, 7, 8, 9, and 10, ps. 27, 28, 31, and 33, respectively of McClure and Soderstrom (1973) as well as in Barkworth et al. (2007, ps. 17-19 passim).�This morphological arrangement (and the general habit it presents) were shown in greater detail in the immediately following slide/caption set.

Leaves seen in these photographs were their natural color (give or take) on the first full day of winter. There were a few senesced (dead at end or completion of their life cycle) leaves on these tillers. Senesced leaves were a light brown or tan in color and appeared clearly in the lower of these two images.

Standard references for the North American�Arundinaria�species (some of which included line drawings and morphological descriptions) were listed in the last caption of this section of Ozark Plateau canebrakes.

Outer bank and first terrace of Elk River, McDonald County, Missouri. Late December; post-peak standing crop, winter semi-dormancy stage.

113. Groups of Ozark bamboo leaves- The leaves on tillers of giant or river cane frequently grow as groups at the shoot apex or grow close enough to each other on branches as to appear to be growing in groups. Actually, there are typically two up to, perhaps, five or even seven leaves growing ranked or in rank order (alternate leaf arrangement) per branch, but there is secondary branching off of the primary branch--sometimes as many as three primary branchees in one unit--that originates from the culm of the shoot, specifically a tiller�(see figures 7, 8, 9, and 10, ps. 27, 28, 31, and 33, respectively of McClure and Soderstrom, [1973]).�In these figures (extraordinary line drawings by botanical illustrator Elmer Smith),�McClure and Soderstrom (1973) distinguished young versus mature shoots and mid-shoot from leafy shoot tips�(known as topknots) of�A. gigantea�ssp.�gigantea. In the leafy shoot tip (apex of the shoot) there can be both leafy and leafless flowering branches (these flowering shoots being very infrequent in production) with upwards of four to six branches at this shoot apex (figure 10, p. 33,�McClure and Soderstrom [1973]).�Some or most of these branches can develop topknots�or, alternatively (and infrequently), inflorescences (usually interpreted as panicles). No spent (dead) inflorescences or spikelets were found in canebrakes shown here.

In the morphological fashion of this branching pattern just described, upper portions of bamboo shoots have a bushy appearance. A very distinctive leaf arrangement in�Arundinaria species is the�topknot, a cluster of four to, say, seven leaves growing in a fan-like arrangement or digitate pattern much like that of tail feathers in birds. This habit can be seen in these two slides as well as was the two slides in the immediately preceding slide/caption set, the second of which provided details of the topknot. Advance to the next slide please ...

Outer bank and first terrace of Elk River, McDonald County, Missouri. Late December; post-peak standing crop, winter semi-dormancy stage.

114. Giant or river topknot- Two views of the fan-shaped cluster of leaves or topknot�(Barkworth et al., p.17-18) at the apex of tillers of giant or river cane growing in a local canebrake on the outer bank of Elk River in the western Springfield Plateau in southwest Missouri. The bamboos (subfamily, Bambusoideae) are the only grasses in North America that have theleaf cluster or topknot. It is also probably a nutritional fact--although it cannot be shown axiomatically like the fact of leaf clusters or topknots--that river or giant cane is the single most palatable grass--native or introduced--growing in the Ozark Plateau and the state of Missouri. Unfortunately, this palatability works agains this marvelous grass as livestock so relish its leaves and young shoots that even in vast canebrakes it is readily "grazed-out" (eliminated by overgrazing).�Leithead (1971, 1976, ps. 50-51)�provided very useful notes on management of this invaluable, native, woody grass.

The�topknots�on tillers in this canebrake typically consisted of four to seven leaves with five or six being the most common number of leaves. This was consistent with line drawings presented in McClure and Soderstrom (1973, figs. 7, 8, 9, and 10, ps. 27, 28, 31, and 33, respectively) as well as less-detailed works like Hitchcock and Chase (1951, fig. 1, p. 28), Barkworth et al. ((2007, ps. 19), and Shaw (2012, p. 237).�Leaves�of giant cane do have a minature (almost rudimentary) petiole that is officially designatedas a pseudopetiole (refer to Barkworth et al. [2007, ps. 17-19]).

Standard references for river or giant cane include all the usual suspects(an expression from the classic picture-show, Cassa Blanca, including Silveus (1933, ps. 1-4), Fernald (1950, p. 96), Hitchcock and Chase (195, 1ps. 27-29), Steyermark (1963, p. 81), Kucera (1961, ps. 14-15; 1998, ps. 25-26 ),�Leithead (1971, 1976, ps. 50-51), Gould (1975, p. 40), Judziewicz et al. (1999,passim, esp. ps.194-198), Barkworth et al. (2007, ps. 17-19), Tyrl et al. (2008, ps. 52-53), and Shaw (2012, ps. 236-237).�Less known references included Judziewicz et al. (1999) and "in-a-league-by-itself" treatment was that of the bamboo giantMcClure, and aided by Soderstrom, (1973, ps. 23-46) which remains the most comprehensive, detailed treatment bar none of Arundinaria gigantea, including its various subspecies.�(Soderstrom edited the almost-readypublication of the recently deceased McClure; extraordinary line drawings were done by botanical illustrtor E. Smith). Barkworth et al. (2007, ps. 17-19) did not recognize subspecies of �A. arundinaria, but their treatment was condensed and elementary (and their line drawings generic and indefinite) as compared to those of �McClure and Soderstrom, (1973, ps. 23-46).

Leaves in these slides appeared more-or-less in their their natural color as photographed on the first full day of winter right after arrival of a "blue norther", a cold Arctic air mass.

Outer bank and first terrace of Elk River, McDonald County, Missouri. Late December; post-peak standing crop, winter semi-dormancy stage.

Common Reed: Reedbank or Reedbed (Carrizal, Spanish for Reedbed)

Some of the greatest diversity-- from standpoint of both species and plant community-- in range vegetation occurs in wetlands. Wetlands are also extremely diverse including such important range plant communities as those of riparain, marshes, wet prairies, swamps, estuaries, wet mountain meadows, and playa lakes. Classification and delineation of wetlands has traditionally varied almost as much as wetlands themselves depending on criteria used and objective (eg. legal, historical, ecological, etc.). Even from these more specific perspectives or discipline emphases the bases for classification or recognition frequently are different. For instance ecological distinction and description has been varied depending on criteria such as biotic community, vegetation, body of water features, soil properties, or duration of water coverage on land surface. In this regard the outstanding encyclopedia, Wetlands, by Mitsch and Gosselink (2000) is definitive (for eg. see Chapter 21 for wetland classification).

One of the wetlands featured variously in the current publication was that of streams (and associated landscapes) because the vegetation of such included several range (rangeland and forest) cover types. Vegetation of the the Rio Grande provided examples of willow scrub and mesquite bosque that were treated under Miscellaneous Shrublands. A third major range plant community of the Rio Grande was patches of grassland dominated by or exclusive communities of common reed or, in Spanish, carrizo (Phragamites communis= P. australis). Such "reedbanks" or "reedbeds"(the Spanish of which is "carrizal"), as they are commonly known, were covered here because they are a minor (very small in area) yet unique and very important grassland that can be interpreted as a range cover type. Reedbanks are a kind of grassland wetland comprised primarily of a native North American grass species that forms or is part of natural range vegetation. This range vegetation was viewed herein as climax.

This was consistent with its classification or designation as the Phragmites communis-Carex spp, et al. Series which together with the Distichlis stricta-Allenrolpea sp Series made up the Chihuahuan Interior Marshland (Brown, 1994, Fig 104, p. 177). Unfortunately, Brown et al. (1998, p. 46) erroneously omitted the Phragmites-Carex Series (a biotic community) from Chihuahuan Interior Marshland, 243.6. The Saltgrass Series, 243.61, included in Brown (1994, Fig , p. 177), was given in Brown et al. (1998, p. 46); therefore, the reedbed vegetation should be Common Reed or Carrizo (Brown , 1994, p. 336)-Sedge Series 243.62.

Common reed was long recognized as forming or developing into the reed-swamp association (Warming, 1909, p. 145; Tansley, 1923, ps. 31; Clements, 1936, ps. 264, 278; Polunin, 1960, p. 463). In fact, as was detailed in the Literature Review (Associations and Consociations section of Range Type), Phragmites communis was one of the most commonly used examples of association (or associes, the seral counterpart) in the Anglo-American School of Plant Ecology.Clements (1920, p. 110) reed-swamp was "[t]he most familar example" of an associes that was usually composed of three consocies one of which was P. communis. Consocies was Clements' term for the seral counterpart of consociation.Common reed (as the "reed-swamp") was one of the "kinds" of vegetation used in derivation of "type", including cover or dominance types (thus, rangeland and forest cover types) from the plant association (again, see Literature Review).

Inclusion of common reed (forming reedbeds or reedbanks) as the common reed cover type provided photographs of vegetation that was of historic significance in development of the cover type concept. Reedbanks were also instructive of a very restricted but a highly productive kind of rangeland and critical habitat along an important stream in the Trans-Pecos Basin and Range. The reedbank or reedbed provided an example of range vegetation along a stream inside a general desert and desert climate. Like willow scrub and mesquite bosque, reedbank was not desert vegetation and therefore these was not included with the Chihuahuan Desert but rather were interpreted as miscellaneous forms of shrubland and grassland, respectively.

116. Route of the river- Land forms and vegetation defined by two rivers:. the Big Bend stretch of Rio Grande in the Trans-Pecos drainage of the Basin and Range province provides an environment for riverine vegetation in the midst of the Chihuahuan Desert. In the far background the Rio Grande emerges from Santa Elena Canyon. This is a chasm with sheer walls up to 1500 feet deep cut through the limestone of Sierra Ponce (the Chihuahua side in the left two-thirds of the background) and Mesa de Anguila (the Texas side in the right one-third background) by sand-laden, moving water over geologic time. Sierra Ponce- Mesa de Anguila are a mesa, "an uplifted block of Lower Cretaceous limestone", that is the remainder of a former and much higher mesa from which the softer Upper Cretaceous limestones and shales were eroded (Maxwell, 1968, ps. 87-89). Even short of its previous glory the mesa is a backdrop land form to a once-mightier river that now relies on another river, the Rio Conco, for most of its water.Even with most Rio Grande water dammed and diverted for human agricultural and municipal uses, there is adequate aquatic environment for well-defined range cover types.

Immediately along the river banks (or back a short distance where there are recently formed sand bars) there is a zone of riparian vegetation made of two end-to-end disparate plant communities or, perhaps more descriptively, stands: 1) willow and 2) reed. The woody vegetation (a willow shrubland) and the herbaceous vegetation (a grassland of common reed [Phragmites communis]) are each spatially short and narrow "strips" of distinct plant communities which obviously interact yet are physically segregated. There are of course other plant species in the willow and reed communities but the two species so dominate their respective vegetation that most other species seem of miniscule importnce, at least from standpoint of range cover types.

Adjacent to riparian vegetation on the river flood plain and covering a much broader breath of rangeland is vegetation known as bosque or bosques (mostly an Anglo-adopted Spanish word). Bosque vegetation is dominated by mesquite and other woody plants, most of which are of shrub rather than tree form and dimension.

Willows are also usually shrubs or, at most, small trees. In restricted locations along short stretches of the Rio Grande riparian vegetation consist of large cottonwood trees, often with willows as a lower woody layer, that form small gallery forests. These gallery forests were treated separately under Miscellaneous Forest Types and Southern and Central Forests (Woodlands and Forests). Gallery forests were not discernable in these landscape-scale photographs.

Technical Note: The term riparian may have somewhat different meanings or connotations as used Hydrology, Aquatic and Riparian Ecology, Range Management, or Forestry not to mention legal usages in the extremely specialized area of water law and policy. The Glossary of Hydrology (Wilson and Moore, 1998) defined riparian thusly: "Pertaining to or situated on the bank of a body of water, esp. of a watercourse such as a river; e.g. 'riparian land' situated along or abutting upon a stream bank..." (listed as synonyms were riparial and riparious). A Glossary of Terms Used in Range Management (Jacoby, 1989), an official publication of the Society for Range Management, defined riparian as: "Referring to or relating to areas adjacent to water or influencecd by free water associated with streams or rivers on geologic surfaces occupying the lowest position on a waatershed". Jacoby (1989) defined riparian zone: "The banks and adjaacent areas of water bodies, water courses, seeps and springs whose waters provide soil moisture sufficiently in excess of that otherwise available locally so as to provide a more moist habitat that that of contiguous flood plains and uplands".

Clearly, hydrophytic or streamside (= riparian) vegetation such as that dominated by willows and reeds on the edge of the Rio Grande constitutes plant communities not only different from other vegetation, but growing on a different environment from that on and of the flood plain. Alternating patches or strips of willow and reed vegetation were clearly visible in both of these photographs. Also readily discernable was the more extensive (and drier-soil) area of mesquite bosque that had developed adjacent to riparian zone vegetation.

Land in the immediate foreground was on an upland overlooking the Rio Grande and Sierra Ponce-Mesa de Anguila that provided a vista of riparian and flood plain vegetation. Vegetation on the upland foreground (vista point) was Chihuahuan Desert dominated by the climax species, creosotebush (Larrea tridentata) and tasajillo or pencil cholla (Opuntia leptocaulis). Quite a study of and contrast in range vegetation.

Big Bend National Park, Brewster County, Texas. June.

117. Two Rio Grande riparian range types- On the north bank of the Rio Grande (south bank is the Rio Bravo) the two major and segretated riparian plant communities of this strean were holding forth: 1) willow scrub made up primarily of sandbar or coyote willow (mostly Salix interior= S. exigua) on the left and 2) common reed grassland on the right.

The back "curtain" of this study in range vegetation was the immense block of Lower Cretaceous limestone that formed the mesa, Sierra Ponce, in Chihuahua. The formation outcrops from top to bottom were: Santa Elena, Sue Peaks, and Del Carmen (Maxwell, 1968, Fig. 92, p. 89). This stretch of river was just east of Santa Elena Canyon.

Rio Grande, Big Bend National Park, Brewster County, Texas. June, early estival aspect and pre-bloom stage.

Directional Note: The Rio Grande willow scrub and mesquite bosque range types were treated in the Rio Grande (Rio Bravo) Vegetation under Miscellaneous Scrub Types (Shrublands).

118. Santa Elena carrizo- Stand of common reed (carrizo in Spanish) at mouth of Santa Elena Canyon. An example of reedbed or reedbank vegetation. Common reed forms dense colonies that develop as linear strips of single-species vegetation which alternate with willow scrub communities of similar size and shape as the two major types of riparian vegetation growing along the Rio Grande as it flows through the arid climate of the Chihuahuan Region, including the Chihuahuan Desert, and eastward into semiarid climate where the regional vegetation is a mixed grass species-thornscrub savanna.

Common reed (Arundineae tribe of Arundionoideae subfamily) is one of the most widely distributed grass species in the world growing in North America, South America, Africa, and Eurasia. Polunin (1960, p. 98) reported that common reed "is often claimed to be the most widely distributed vascular plant species in the world".

Carrizo was the native grass most commonly used for purposes aside from forage or game habitat by Indians and, later, Mexican peasants living along Chihuahuan Region rivers like the Rio Grande. Maxwell (1968, ps. 105-106) described use of the culms of common reed as building material for huts. Arrow shafts were supposedly fashioned from the smaller distal portions of carrizo culms.

Santa Elena Canyon, Big Bend National Park, Brewster County, Texas. June, late pre-bloom stage. No FRES, Kuchler (1964, in Garrison et al., 1977), or Society for Range Management (Shiflet, 1994) units for common reed or carrizo range cover type. Erroneously omitted by Brown et al. (1998, p. 46) but there should be a Common Reed-Sedge Series, 243.62, of Chihuahuan Interior Marshland, 243.6. Chihuahuan Deserts- Low Mountains and Bajadas Ecoregion, 24c (Griffith et al., 2004).

119. Reeds along the Rio- In this short reach of the Rio Grande (Rio Bravo) common reed or carrizo was growing on both stream banks in direct contact with river water. Reedbeds or reedbanks were the riparian vegetation. This riparian strip of grassland was interrupted at intervals by willow scrub (or, alternatively, riparian willow vegetation was "broken" periodically by stands of carrizo). Though extremely small in size (narrow often short colonies) reedbeds were the major source of forage along the river. Reedbanks appeared to produce appreciably more biomass than willows, but a combination of reed forage and willow browse provided a good combination diet plus cover for game animals like deer. Water, high-quality feed, and shade makes for great wildlife habitat. Reedbeds were ideal range. Maxwell (1968, p. 105) told how Mexican stockmen fired the carrizal (reedbeds) in the winter to insure green feed from tender carrizo shoots in the spring.

Common reed was far more valuable than would be suspected by the extremely small portion it made up of total Chihuahuan Region vegetation. This was yet another example of the incaluable importance of riparian range vegetation. Importance of reedbanks in prevention or reduction of soil erosion was shown in the next two photographs.

Rio Grande (Rio Bravo), Big Bend National Park, Brewster County, Texas. Sierra Ponce in Chihuahua in background. June, early estival aspect and late pre-bloom stage. NO FRES, Kuchler (1964, in Garrison et al., 1977), or Society for Range Management (Shiflet, 1994) units for carrizo or reed rangeland cover type. Should be a Common Reed-Sedge Series, 243.62, of Chihuahuan Interior Marshland, 243.6 (Brown et al., 1998, p. 46). Chihuahuan Deserts- Low Mountains and Bajadas Ecoregion, 24c (Griffith et al., 2004).

120. Reedbed on the river bank- Common reed or carrizo growing on bank of the Rio Grande. Photographs showed structure typical of a stand or colony of carrizo. Specifics of bank-cutting was unknown. Obviously there had been recent flooding without enough subsequent river flow to carry away shoots of reed. Whether this was natural (= geologic) or accelerated soil erosion was not known. River banks are dynamic with some stream banks always being eroded while other banks are being rebuilt (vertical accretion) by sediment deposited from eroded banks. Soil (mostly sand) eroded from this bank was being deposited as a point bar on a stream meander downriver. This location was in close proximity to mouth of Sant Elena Canyon where confined stream flow rushed onto a broad floodplain of that reach of the Rio Grnde. A handy reference or guide for discussion of stream processes and patterns was the small book by Morisawa (1968).

Big Bend National Park, Brewster County, Texas. June, late pre-bloom phenological stage. No FRES, Kuchler (1964, in Garrison et al., 1977), or Society for Range Management (Shiflet, 1994) for reedbed (carrizal) range cover type. If had not been mistakenly omitted from Brown et al. (1998, p. 46) would be Common Reed-Sedge Series, 243.62, of Chihuahuan Interior Marshland, 243.6. Chihuahuan Deserts- Low Mountains and Bajadas Ecoregion, 24c (Griffith et al., 2004).

121. Reeds in the Basin and Range- Common reed or carrizo on a bank of the Rio Grande. This photograph of the north bank of the Rio Grande with Bofecillos Mountains in the background showed physiography of the Basin and Range province from a "close-in" perspective. This slide also presented a good example of the physiogonomy of a reedbed as riparian range vegetation. The perennial nature of common reed was apparent, including some perennality in shoots. More-or-less dead shoots from previous growing seasons (those of last year or several years ago was not determined) were mixed in with current season's shoots. Some of last year's shoots had grown small secondary shoots from intercalary meristem of internodes. Thus plants of common reed sometimes have perennial shoots (ie. in this herbaceous species it is not just the rootcrown or proaxis that lives for more than one growing season; some shoots of this herbacsous species develop lateral shoots proving that there is some aboveground perennating tissue).

Presidio County, Texas. June, early estival aspect and late pre-bloom stage. No FRES, Kuchler (1964, in Garrison et al., 1977), or Society for Range Management (Shiflet, 1994) designations for reedbed (carrizal) rangeland cover type. Common Reed-Sedge Series, 243.61, of Chihuahuan Interior Marshland, 243.6 (Brown et al., 1998, p. 46, but inadvertently left out).Chihuahuan Deserts- Low Mountains and Bajadas Ecoregion, 24c (Griffith et al., 2004).

122. "Reed swamp" on the Rio Grande- It was explained in the introduction to the reedbed portion that one of the most commonly used examples of a plant association or, the seral equivalent, associes by pioneer British and American plant ecologists (eg. Warming, Clements, Weaver, Tansley) was the so-called "reed-swamp". This included the Phragmites comminus or P. australis consociation or, the seral unit, consocies. Since those fledging years in which successional and vegeetation classification concepts were formulated wetland specialists, including ecologists, have defined swamp as wetland vegetation dominated by woody plants whereas wetland plant communities dominated by herbaceous species are marshes. Common reed-dominated vegetation (which is limited to permanently wet or, at least, very moist soils) is now recognized as a marsh not swamp cover type. Even the term marsh (ie. reed marsh) is not as appropriate or as descriptive of the Rio Grande reedbeds as the title, common reed-dominated riparian vegetation. Plant communities that are restricted to stream banks or adjacent habitats should most properly be entitled and described as having developed in the riparian zone; hence,as riparian vegetation.

Reedbanks are, therefore, riparain range plant communities or, in terminology and context of this publication, the common reed (Spanish, carrizo) or reedbed (Spanish, carrizal) range cover type is riparian range. Plant communities of P. australis in many other areas, including elsewhere in North America, are actual or true marshes. These are properly termed reed marshes. One of the largest and most notable of these reed marshes is the vast network of P. australis and Typha spp.- dominated wetlands known as the Mesopotamian marshlands that developed over the immense lowlands of the Tigris and Euphrates Rivers. Most of this reed-cattail marsh was in Iraq and home to the Ma'dan or Marsh Arabs. This wetland ecosystem and the Ma'dan cultural system is slowly being restored following occupation by American military forces in the current war for Iraqui "liberation". Compared to such vast reed marshes the scattered patches of riparian reedbed range along banks of the Rio Grande were mere parcels of a riparian oasis in the Chihuahuan Desertscrub and Tamaulipan thornscrub savanna. Common reed range communities were nontheless priceless for their forage, cover, soil protection, biodiversity, and so on.

This view of a reedbed was on the outermost edge of the Rio Grande where young shoots of carrizo were in early growth. Older (larger) shoots, including both dead and previous-growing season shoots having intercalary branches as well as current- season shoots, were farther back (deeper) in the colony. Most grazing by ungulates would be on this vegetational "frontier" with consumption of biomass deep inside the reedbed limited to smaller vertebrates like rodents and invertebrates (eg. insects).

Big Bend National Park, Brewster County, Texas. June, early vegetative growth stage (see "very next" photograph). No FRES, Kuchler, Society for Range Management (Shiflet, 1994) units for what was obviously a reed rangeland cover type. Common Reed-Sedge Series, 243.62, of Chihuahuan Interior Marshland, 243.6 (Brown et al., 1998, p. 46 but inadvertently omitted). Chihuahuan Deserts- Low Mountains and Bajadas Ecoregion, 24c (Griffith et al., 2004).

123. Reed reproduction- Asexual reproduction in common reed or carrizo includes all three types of shoots in grasses: 1) tillers which are vertical, intravaginated shoots, 2) rhizomes which are horizonal, belowground, extravaginated shoots, and 3) stolons which are horizonal, aboveground, extravaginated shoots. Seen here were stolons that had produced tillers at the "frontier line" of the advancing colony of this clonal or modular plant.

At edge of water line in bed of Rio Grande, Presidio County, Texas. June, early growth phenological stage.

124. More reed reproduction ("if and when it rains")- Two stolons developed from a rhizome of common reed growing beneath sand along the Rio Grande. This soil was on the backside of a reedbed (the dry side or side of the reed colony away from the water's edge). Green leaves had recently emerged from these shoots which otherwise appeared dead. Will the new shoots (young ramets of the established genet) survive by being able to translocate water, mineral nutrients, and photosynthate from the parent plant or are they dependent on water filling the pores of the dry sand in which they were rooted in?

Presidio County, Texas. June, early summer but early "green-up" for these young 'uns.

125. Reed tiller tops- Distal or apical portions of tillers of common reed or carrizo. The most obvious shoots of common reed or carrizo are tillers, but these tall vertical.shoots typically arise from rhizomes or stolons the "unsung shoots". Polunin (1960, p.98) felt that P. communis (= P. australis) was not only one of the most widely distributed plants on Earth but also one best adapted for dispersal as it has wind-carried, parachute caryopses; water-borne, buoyant rhizomes; and soil surface-covering stolons. Dispersal of propagules or disseminules by wind is anemochory while dispersal of germules by water is hydrochory, and dispersal by gravity is barochory. Dispersal of propagules by more than one agent or mechanism is polychory. Is it any wonder that the polychorous P. communis has one of the largest discontinuous (= disjunct) ranges (geographical distributions) of any extant plant species?

Chapman (1996, p. 145) summarized sexual reproduction in P. australis. He reported that establishment of new reed plants from seed varied tremendously. Although common reed flowers plenteously, grain production is generally low and seedling establishmen is usually rare in native populations. Sexual reproduction is much greater in domestic reed than in natural reedbeds (Chapman, 1996, p. 146). Even phenotypic variation within a clone (genet) is fairly limited.

It would be interesting to know if the isolated stands or colonies of carrizo along the Rio Grande are distinct genotypes (genets, clones, or modules) or if some of these colonies are of the same clone with a "broken" distribution due to disturbances (possibly flood, fire, grazing, human harvest for building materials) that resulted in death of parts of clones (loss of ramets) at various intervals in space and time.

Tillers in the upper (top) photograph were young, current-season's shoots. These tillers had arisen in the current growing season from rhizomes. There were no secondary shoot or "branches" arising from intecalary meristem along the tiller. Tillers in the lower (bottom) photograph were last year's or the previous growing season's shoots with secondary tillers or "branches" that developed from intecalary meristem during the current growing season. The "branches" or secondary (intecalary) shoots (which are also tillers) demonstrated that some tillers are perennial. It is not just the plant (the genetic individual= genotype= genet) that is perennial, but some shoots (some modules= clones) are perennating organs.

Tillers in both photographs were on the river (water) side of reedbeds growing along the Rio Grande. Presidio County, Texas. Advanced pre-bloom phenological stage.

126. Reed flowers- Infloresecence of common reed or carrizo at early anthesis. This speccimen was growing on a freshwater marsh on South Padre Island, Cameron County, Texas.

127. Reed spikelets- Portion of panicle of common reed or Rouseau cane featuring spikelets. Cameron County, Teaxas. February, seed ripe phenological stage.

128. Riverbottom vegetation without a place to go- Along a recently flooded Little Missouri River there were two distinct range plant communities: 1) a reedbed of common reed (Phragmites communis= P. australis), the vegetation zone nearest the river comprised solely of short, green grass and 2) a floodplain forest co-dominated by eastern cottonwood (Populus deltoides subsp. monilifera) and peachleaf willow (Salix amygdaloides) adjoining the reedbed.

The stump in foreground of the second slide was that of a cottonwood that had been felled by feeding beaver (Castor canadensis) with the log (trunk) carried downstream by the recent record flood. Details of this coppicing (sprouting) cottonwood were shown in the next subsequent slide.

Clearly a large part of this range vegetation was forest (and this exmple was also included with treatment of cottonwood-willow forest cover type, the reedbed certainly had to be included in this section.

Along Little Missouri River, Theodore Roosevelt National Park (North Unit), McKenzie County, North Dakota. Late June. FRES No. 17 (Elm--Ash-Cottonwood Forest Ecosystem). K-89 (Northern Floodplain Forest). SAF 235 (Cottonwood-Willow). Cottonwood-Willow Series 221.21 in Plains and Great Basin Riparian Deciduous Forest biotic community 222.2 and Plains Interior Marshland 242.3, but no Common Reed Series given of Brown et al. (1998, p.43 ). Northwestern Great Plains- Little Missouri Badlands Ecoregion Ecoregion 43b (Byce et al., Undated).

129. Browsing in a reedbank- On the immediate or near (oputermost) channel of the Little Missouri River an eastern cottonwood felled by beaver was resprouting at the edge of a cotton-willow riparian forest and a bed of common reed that had just emerged from dormancy. This entire riverbottom vegetation was recovering from recent flooding. There were two distinct floodplain plant communities: 1) the bed of common reed and 2) botttomland forest of eastern cottonwood and peachleaf willow.

Clarifying Note: Beginning students could easily confuse common reed (P. communis= P. australis) with giant reed (Arundo donax) from both similar physical appearance and morphology of plants as well as from the common name of "reed". (The Spanish names, carrizo and carrizal, would apply only to the native P. communis and not A. donax this latter of which was introduced and naturalized after establishment of Hispanic culture.) Both of these arundinoid grasses occur in some areas and, to add to confusion, in the same general habitat therein. Both species grow along the Rio Grande: in fact, sometimes side-by-side (Powell, 2000, p. 46). A. donax is an exotic species introduced from the Old World (Hitchcock and Chase, 1951) that appears to have naturalized. In North America spread of A. donax appears to be strictly by asexual reproduction and not by sexual means because viable seed production seldom if ever occurs. Instead dispersal is by fragmentation of rhizomes and/or root systems or, less commonly, by secondary shoot development from detached tillers. This fragmentation is often human propagation (culms of tillers are used for reed instruments, the entire shoot has some agronomic potential as a biomass fuel crop).

A. donax has become a very aggressive, exotic weed in riparian and general aquatic ecosystems. A. donax should not be propagated along rivers. Rather it should be controlled or, better yet, eradicated if possible from natural range ecosystems such as those of the various range types along the Rio Grande.

This author/photographer did not find any A. donax on the Rio Grande. All photographs of reedbeds presented above consistedly strictly of P. communis (or associated native willow scrub or mesquite bosque).

130. Ecological miracle in the canyon- Two conterminous wetland range plant communities that developed on a small, spring-fed stream emanating from cliffs in Wind River Canyon in the Wyoming Basin. The upper, darker-green band of vegetation was basically a colony of the stoloniferous, thicket-forming red-ozier dogwood that grew--almost exclusively--in the riparian zone. There was also some cover of white (or western) virgin's bower in with the red-ozier dogwood. Farther back from this streamside thicket of red-ozier dogwood and western clematis, Utah juniper, skunkbush sumac, western wheatgrass, and bluebunch wheatgrass formed a shrub-grass savanna. (This scrubland or, more accurately, shrub-grass savanna was introduced above and covered in greater detail in the immediately following two-slide-caption unit.)

The local (microsite) common reed-dominated marsh had broadleaf cattail (Typha latifolia) and smooth scouring rush (Equisetum laevigatum) as associates to common reed. Immediately beyond the reed-cattail-scouring rush marsh (yet still in moist soil) there was a zone of two, naturalized, Eurasian, perennial grasses, Kentucky bluegrass (Poa pratensis) and orchardgrass (Dactylis glomerata). These two cool-season (festucoid), agronomic grasses attested to the mesic zone just beyond the hydric (wet) zone of the marsh.

It was explained variously throughout the Grassland chapter entitled , Meadows and Related Marshes that marshes dominated by common reed are some of the most widely dispersed natural, herbaceous wetlands on Earth. In the older literature these graminoid wetlands were often--and incorrectely--called "reed swamps". Such common reed marshes were interpreted as a seral stage along a hydrosere or, perhaps, as subclimax or, even a lower seral stage, serclimax in the Clementsian monoclimax theory (Clements, 1936, p. 264; Weaver and Clements, 1938, ps. 62-63, 81) and as a hydric, edaphic climax in the Tansian polyclimax theory (Tansley, 1926, p. 31, 33, 34, 36, 59-60).

While of wide distribution around the planet, reed marshes--with notable exceptions of vast marshes in the Mediterranean Region--are typically of small size. This does not in any way negate their importance as refugia for both plant and animal species nor minimize their miraculous occurrence in the most unexpected locations and forms of terrain. Reed marshes are literally oases in regional climates that are substantially drier than the hydric habitat of marshes.

The larger, taller amber-colored stalks were last year's shoots of common reed. The color distinction between these dead stalks and current year's live shoots was obviously more pronounced in the second and third of these three slides which were taken from more of a sideview. By contrast, the first slide was taken from more of a topdown view and at an oblique angle. From that vantage perspective the green of living shoots "outshone" the amber of dead shoots.

Wind River Canyon. Hot Springs County, Wyoming. Late June. For the red-ozier dogwood-dominated riparian vegetation: No FRES, Kuchler unit, or SRM or SAF cover type. Willow-Dogwood Series 232.51, Rocky Mountain Swamp and Riparian Scrub 232.5 of Cold Temperate Swamp and Riparian Scrub 232 (Brown et al., 1998, p. 38). For the common reed-dominated marsh: No FRES, Kuchler unit, or SRM or SAF cover type. In Brown et al. (1998, p. 45) this natural plant community should be a Giant Reed Series, say, 242.83 (that Brown et al. ][998, p. 39] gave for Tropical-Subtropical Marshland and did not give for this unit), in Rocky Mountain Montane Marshland 242. 8 in cold Temperate Marshland 242. Wyoming Basin- Foothill Shrublands and Low Mountains, Ecoregion 18d.

Florida Grasslands

131. Firing a wiregrass or threeawn (Aristida spp.) range in Florida- Use of prescribed fire to control saw palmetto on a Florida wiregrass range. Fire has traditionally been an integral part of Forestry and Range Management in much of the Southeast . This goes back to the American Indians who with their own form of education and training were sound range managers. Avon Park, Florida. February (demonstration burn: too late for proper prescribed fire because grass is too far advanced in its annual growth cycle, but fire did reduce palmetto cover).

132. Saw palmetto (Serenoa repens) set back by prescribed burning- Saw palmetto is distinguished from palmetto (Sabal minor) by presence of aboveground horizontal shoots in the former versus belowground rootstocks (rhizomes) in the latter. Saw palmetto was only top-killed, perhaps not even that, on a Florida bluestem range, but cover was reduced enough to give the greening forbs and native bluestems a competitive advantage. This is the essence of noxious range control. Burning , like anything else, can be overdone. Most desirable grasses are adversely affected by annual burning so this should be avoided. Avon Park, Florida. February.

133. Detail of fire damage- Closer views of saw palmetto top-killed by use of prescribed burning on the Florida bluestem range shown above. Avon Park, Florida. February.

134. Saw palmetto (Serenoa repens)- Saw palmetto on a mixed bluestem Florida range. Various species of bluestems that were growing on this same range were presented below. Avon Park, Florida. February.

135. The ranges of the Southeast, including peninsular Florida, divide at broadest scale into either: 1) wiregrass or threeawn (Aristida spp.) or 2) bluestem (dominance by Andropogon spp) grasslands or forest understories. The bluestem ranges are generally superior from standpoint of both quantity and quality of forage (though as always this varies with and depends on such factors as species, stage of maturity, season, soil fertility, and weathering). Creeping bluestem (Andropogon stolonifer) is one of the major forage grasses in the pine-hardwoods region. It is a climax dominant and major forage producer. It grows on various sites ranging from sandhills to flatwoods and holds its relatively high nutritive value through winter. It is taken readily by range animals yearlong. Creeping bluestem benefits quickly from control of saw palmetto (Leithead et al., 1971). Avon Park, Florida. February.

136. Chalky bluestem (A. capillipes) on a flat woods range site- The flat woods is one of the most common range sites in Florida and chalky bluestem is one of the dominant decreasers on it. The range of this species extends from North Carolina to east Texas. Like creeping bluestem, chalky bluestem produces good forage (including in winter). When managed properly it recovers quickly on abused ranges and even go-back land. Thus it is important in erosion control (Leithead et al., 1971). Avon Park, Florida. February.

137. Detail of the chalky bluestem in the preceding slide.

138. Freshwater marsh of maidencane (Panicum hemitomon)- This consociation of a hydrophytic panicgrass is a productive range site that is usually local in scale but it has a geographic range from east Texas to Florida northward to New Jersey. This is the Maidencane Marsh variant of SRM 819 (Freshwater Marsh and Ponds). It is one of eight major types of freshwater marshes in Florida (Shiflet, 1994). FRES No. 41 (Wet Grasslands Ecosystem). Minor component of K-83 (Everglades).Avon Park Bombing and Gunnery Range, Florida. February, hiemal aspect.

139. Cut-Throat Seep- Like the preceding vegetation, the cut-throat seep is a range site and not a range type per se. It is an edaphic form of the semitropical grassland-savanna complex of southeastern North America, perhaps most widely distributed and extensive in Florida. Cut-throat seeps are one of the better examples of these tropical-semitropical savannas of grassland with their major tree and shrub components. In the continuum of soil moisture gradients--chresard is the classic Clementsian term for the quantity of soil water available for plant use--cut throat seeps are essentially hydric and next to marshes in this context. Soils are typically wet year-round and support cut-throat panicgrass (Panicum abscissum) as a dominant species along with an array of panicoid grasses from maidencane to creeping and chalky bluestems. The widely distributed wax myrtle dominates the shrub layer that on better-drained sites is made up largely of saw palmetto. Slash pine is the major component of the tree layer of this community as on adjoining range sites which demonstrates adaptation of this valuable species to an array of soil moisture regimes. Slash pine does best on lower sites having more available moisture as seen here.

Avon Park Bombing and Gunnery Range, Florida. FRES No. 41 (Wet Grasslands Ecosystem). Variant or phase of K-71 (Palmetto Prairie). SRM 813 ((Cutthroat Seeps). February, hiemal aspect.

Northern Oregon Headlands

There are some range plant communities or even locally important range cover types that defy obvious classification and that do not seem to fit conveniently anyplace. These are a miscellaneous category of vegetation as it were. This can even be the case when grouping into the largest plant community level of biome. One such example is vegetation of beaches or dunes. These sometimes fit readily into the grassland or forest biome or into the more diverse formation of shrubland or scrub. Perhaps grassland and scrub of beach or dune form a savanna that while not meeting the biome distinction nonetheless fit readily into a general and long-recognized category of vegetation (including climax vegetation).

One example is the beach-dune vegetation of certain headland range communities along the Pacific Ocean such as some of those in the northern Coast Range. These natural communities were described by Franklin and Dyrness (1973, ps. 291-300). Beach and dune vegetation of California was discussed generally in Barbour and Major (1995, ps. 223-261).

Examples of beach and dune vegetation along the Coast Range of northern California were presented in the current publication under: 1) Grasslands, California Grasslands and 2) Shrublands, California Chaparral under special heading Pacific Coast Scrub.

The range vegetation presented immediately below was dominated (on other than microsites) by herbaceous species, including pteridophytes (vascular cryptograms) as well as other forbs along with grasses and grasslike plants. This vegetation was clearly not grassland but it fit that biome grouping better than others and was included here in the section or chapter, Miscellaneous Grassland.

140. Range vegetation on stabilized sand dune along Pacific Ocean- The unusual plant community on this high dune was in the Coast Ranges physiographic and geologic province in northern Oregon. This ocean-front and its herbaceous community could be readily visualized and interpreted as a "headland" though certainly not the more characteristic high and rocky headlands found in the same area. The ocean-most extremity of vegetation (foreground) was within the swash zone at high tide such that the beach-bordering vegetation of this range ecosystem could also be viewed as a dune plant community.

Zonation of plant species and groups was remarkably well-defined. The first zone or swash zone (foreground) was composed of salt rush (Juncus lesueurii= J. balticus var. lesueurii). Immediately behind the salt rush zone was another zone dominated by the native legume, giant vetch (Vicia gigantea). In turn the umbels sea-watch (Angelica lucida) and beach lovage (Ligustichum scoticum) formed a small "patch" behind the giant vetch-dominated "strip". Behind the umbel "patch" vegetation was dominated by giant horsetail (Equisetum telmateia) but with the grasses Idaho bentgrass (Agrostis idahoensis), seashore bluegrass (Poa macrantha), and Pacific reedgrass (Calamagrostis nutkaensis) and silverweed (Potentilla anserina).

Yaquina Bay State Park, Lincoln County, Oregon. June. Coast Range- Coastal Lowlands Ecoregion, 1a (Thorson et al., 2003).

141. High sand dune Pacific Ocean range community- This high sand dune immediately beyond the Pacific Ocean beach had been stabilized by a plant community dominated by giant horsetail with several grass species, including seashore bluegrass, Pacific reedgrass, and Idaho bentgrass, serving as associates. Also present, though widely scattered, was Oregon Iris (Iris tenax).

Yaquina Bay State Park, Lincoln County, Oregon. June. Coast Range- Coastal Lowlands Ecoregion, 1a (Thorson et al., 2003).

142. Sand dune range plant community along Pacific Ocean- Adjacent to swash zone of Pacific Ocean in the Oregon Coast Ranges was this natural vegetation of unusual composition. The low shrub in immediate foreground was deer brush (Ceanothus intergerrimus). Behind this shrub was a zone of vegetation consisting of giant horsetail, Pacific reedgrass, seashore bluegrass, Oregon iris, giant vetch, and silverweed.

Yaquina Bay State Park, Lincoln County, Oregon. June. Coast Range- Coastal Lowlands Ecoregion, 1a (Thorson et al., 2003).

143. Salt rush (Juncus lesueuri= J. balticus var. lesueuri)- A dominant range species at the edge of swash zone that formed the lowermost segment of vegetation at base of sand dunes along the Pacific Ocean in the Oregon Coast Ranges. Inflorescences at anthesis.

Yaquina Bay State Park, Lincoln County, Oregon. June.

144. Oregon iris (Iris tenax)- One of the conspicuous range forbs in a ocean-front sand dune herbaceous community.Yaquina Bay State Park, Lincoln County, Oregon. June.

Basin or giant wildrye (Elymus cinereus) range type

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

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

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

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

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

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

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

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

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

146. Basin wildrye range- Interior of the relict Great Basin wildrye grassland presented in the immediately preceding photograph showing species composition, plant morphology and dispersion, and community structure. Botanical composition and vegetation stucture were simple: one species and one layer of vegetation. Plant growth form was largely cespitose (=bunched or clumped). Basin wildrye does have short rhizomes, but it retains the basic morphological form of a bunchgrass.

In the virgin landscape much of the graminoid vegetation here in the Silvies River valley was wetland (bulrush-dominated marsh) and associated mesic grassland consisting largely of basin wildrye. Much of the marsh was converted to hay fields of introduced Eurasian perennial grasses while the climax virgin grassland was farmed, overgrazed, and otherwise largely destroyed or highly modified so that most of the basin wildrye was replaced by other plants.

Unfortunately the value of basin wildrye communities was not appreciated until most of it was gone. Even plant ecologists seem not to have paid much attention to this robust, highly productive species and the range plant communities dominated by or made up of it (see for eg. brief coverage given by Franklin and Dyrness, 1973, p. ). Even the Society for Range Management (Shiflet, 1994) did not designate a basin wild rye rangeland cover type. Perhaps so much of the original basin wildrye vegetation was gone that it had little practical use.

Some taxonomic authorities recognized two varieties of Elymus cinereus. Hitchcock and Cronquist (1973) distinguished between E. cinereus var. cinereus having glabrous sheaths and E. cinereus var. pubescens having pubescent sheaths.

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

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

Texas Pineywoods Naturalized Woodland/Savannah Pasture

Throughout much of eastern North America the original forest vegetation was highly modified by European (white) man. Over much of this vast domain forest were cleared and converted into cropland, including introduced pasture. Some land was cutover forests that through secondary plant succession partially recovered as forest vegetation, often with appreciable propotions of naturalized introduced plant species. Over the course of time (decades in many instances) much of this land that had been farmed was either abandoned (in most cases of which it reverted to second-growth forests) or used as grazing land managed extensively and, usually, improperly (ie. mostly just grazed and fertilized, reseeded, renovated, etc. only periodically, if that, and generally used without regard to proper husbandry-- of pasture or livestock). Some of the second-growth forest on cutover land was also grazed/browsed by livestock and/or wildlife. Aside from stocking with livestock and supporting free-ranging wildlife that might be hunted, about the only human input to this mostly mismanaged, more-or-less defacto pasture was mowing or shredding (when the weeds and brush had gotten "too thick" and/ or grass shoots had grown so tall as to appear unsightly and prompt an oral response--which might or might not be followed by action--such as "It looks sorta rough; guess I ought to knock her down some").

This category of grazing land is probably the major form of naturalized pasture (naturalized range) in North America. (Naturalized annual grassland in California is the most recognized type of naturalized range, and it is generally much better managed than naturalized pasture to the east.) In northen and central regions of North America naturalized pasture (grazing land and range are appropriate synonyms for pasture when the adjective naturalized is applied) is typically dominated by Eurasian domesticated perennial grass species and/or weedy (ruderal) annual grasses of cool-season tribes especially Festuceae (Poaeae) and Aveneae. In more southern regions of North America naturalized, introduced perennial grasses tend to be of African or South American origin and in warm-season tribes, notably Andropogoneae, Paniceae, and Eragrosteae. Annual grasses on these southern anthropogenic pastures--which can be grasslands, savannahs, woodlands, or forests-- include many of the same genera and species (in particular Bromus, Avena, Hordeum) as those on naturalized grazing lands to the north and east and on the Pacific Slope. Some of the annual cool-season grasses in the south are native. Little barley (Hordeum pusillum) is the most common (widespread) example. In addition to these annual festucoid grasses, annual panicoid species are frequently abundant on naturalized southern range. Some of these are also native. Texas millet or Texas panicgrass (Panicum texanum) is perhaps the most common of these species. Most warm-season annual grasses are naturalized (primarily weeds imported from Eurasia). Hairy crabgrass (Digitaria sanguinalis), southern crabgrass (D. ciliaris), and stinkgrass (Eragrostis cilianensis ) are examples of this group that are common in the Texas Pineywoods.

An example of naturalized pasture typical of southern North America was provided by a Texas Pineywoods woodland (or savannah, depending on interpretation) dominated by native oaks, water oak (Quercus nigra) and southern red oak (Q. falcata), in the tree layer and agronomic (introduced) perennial panicoid grasses, bahiagrass (Paspalum notatum) and Vaseygrass (P. urvillei) in the herbaceous layer. Both of these domestic Paspalum species were introduced from South America. Interestingly enough, Dallisgrass (P. dilatum) which is a more widely naturalized Paspalum species in Texas was not observed on this example pasture by the author. Apparently Dallisgrass is not competitive with the rhizomatous bahiagrass and taller, ranker Vaseygrass in the more mesic habitat of the Pineywoods, at least on sites represented by the example shown here. There were some native Paspalum species, including thinseed paspalum (P. setaceum) and brownseed paspalum (P. plictum), but these were "totally overwhelmed" by the two exotic domestic paspalums. Other grasses present included the natives, purpletop (Tridens flavus), broomsedge (Andropogon virginicus), beaked panicgrass (Panicum anceps), and Texas millet. With the exception of the latter, these native grasses were most common under oak canopy. It seemed that they were more competitive with the two dominant Paspalum species mostly under shade. Common bermudagrass (Cynodon dactylodon) was present, but not abundant. Like naturalized Dallisgrass, bermudagrass appeared to be not competitive with bahiagrass and Vaseygrass. The most common forb in this naturalized pasture was wooly croton or hogwort (Croton capitatus), a native annual.

The herbaceous layer(s) of vegetation of this grass-oak woodland was, once again, largely dominated by introduced species of Paspalum (P. notatum and P. urvillei) as well as such other naturalized exotic grasses as Cyndon dactylon and even less locally common exotics as P. dilatum and Bothriochloa ischaemum along natives like Tridens flavus, Andropogon virginicus, and and scattered individuals of various Paspalum species like P. plicatulum, P. setaceum, P. pubiflorum, and P. bifidum. Herbage was largely or overwhelmingly of naturalized alien panicoid grasses. This range vegetation is not widely distributed like California annual grassland so there was no SRM rangeland cover type for it. Nor was there an SAF forest cover type for woodland dominated by water oak and southern red oak.

148. Naturalized woodland (or savannah) pasture in Texas Pineywoods- An herbaceous understorey dominated by bahiagrass with Vaseygrass as the associate species that developed beneath scattered young trees of water and southern red oak was used as naturalized range for beef cattle and white-tail deer at western edge of Texas Pineywoods. The largely anthropogenic (man-made) vegetation of this woodland range included some native grasses of which the major perennials were (in approximate order of abundance) purpletop, broomsedge bluestem, beaked panicgrass, thinseed paspalum, and brownseed paspalum. These species were most abundant (greatest density with highest cover) in shade of oaks. The native Texas millet was the most common and productive annual grass. The introduced( and widely naturalized) common bermudagrass and Johnsongrass were also locally abundant species, but Dallisgrass was not present at other than trace amounts (ie. it was not found at cover or density that was floristically meaningful). There were some Carex and Cyperus species, but these were infrequent and could not be identified without inflorescences which had been removed by recent mowing. The most common forb was the native annual euphorb, hogwort or wooly croton.

This pasture had been recently destocked of cattle and shredded. Prior to destocking it had been grazed to light degree of use, but subjected to frequent mowing at comparatively low stubbgle height. Aggressiveness and highly competitive features of bahiagrass and Vaseygrass effectively excluded most other grasses. These two agronomic paspalums apparently were less competitive under conditions of shade. It was likely that frequent, short mowing (shredding) had limited cover and density of Johnsongrass, a common and aggressive weedy perennial through much of southern and central North America. The conventional wisdom that Johnsongrass can tolerate almost any abuse except repeated close grazing and/or mowing was probably demonstrated on this naturalized pasture.

Domination of herbaceous cover by bahiagrass became a widespread phenomenon over much of the North American southland within a few decades of its introduction. Bahiagrass became an aggressive weed much like Johnsongrass though it has a considerably smaller area of distribution. Since introduction of bahiagrass numerous cultivars of bermudagrass were developed that are much more productive (higher-yielding) than bahiagrass or common bermudagrass, but bahiagrass and, to a lesser extent, common bermudagrass outcompete hybrid bermudagrasses and take over fields planted to the superior forages (and inferior competitors). The ranker-growing Vaseygrass maintains cover and spreads less effectively than bahiagrass under heavier defoliation, but it typically fares better under closer mowing and grazing than the tall-growing Johnsongrass and the shorter, stoloniferous bermudagrass (at least under the warm, mesic.habitats of the Pineywoods). Although Vaseygrass is semirhizomatous it usually grows more as a bunchgrass (much like the smaller Dallisgrass), which combined with its elevated apical meristems, puts it at competitive disadvantage to bahiagrass.

Freestone County, Texas. October; grain-ripe phenological stage in the two major paspalums. No units of native vegetation were applicable for this naturalized woodland pasture. East Central Texas Plains- Floodplains and Low Terraces Ecoregion, 33f (Griffith et al., 2004).

149. Naturalized grass under native trees- Physiography and structure of a naturalized herbaceous layer beneath scattered young to mid-age water and southern red oaks. Bahiagrass and Vaseygrass were the principal herbaceous species with Vaseygrass the associate to co-dominant species on local spots (microsites) having less cover of bahiagrass which was the overall dominant. Common bermudagrass and Johnsongrass were other locally abundant introduced perennial grasses. Native perennial grasses included (in this approximate order of abundance) purpletop, broomsedge bluestem, beaked panicgrass, thinseed paspalum, and brownseed paspalum. Texas millet was a common and even locally dominant native annual grass. There were some species of caric sedge (Carex spp.) and flat or umbrella sedge (Cyperus spp.) which could not be identified in their existing vegetative state. Hogwort or wooly croton, a native monecious euphorb, was the major forb.

The oak species were present as individuals as well as clumps (assemblages) of trees. Bahiagrass and Vaseygrass were comparatively less abundant under tree shade, sometimes even being associate to mere incidental species under dominance of native grasses. Tree regeneration was limited to seedlings so short that the shredder blades passed over them. Browsing was not evident, probably because mowing had twisted off taller seedlings and saplings. There were some larger (taller) young trees close enough to existing trees that the rotary shredder could not get to them. The history of mowing and grazing management was unknown though this man-modified naturalized pasturage was representative of others in this western edge of Texas Pineywoods.

The spatial distribution of trees on this naturalized grazing land was subject to interpretation as either woodland or savanna. There were stands of water and southern red oak in which tree density and proximity were such that crowns interlocked to form a forest canopy (though a relatively open canopy layer). There were other locations on this naturalized range where there were no trees and enough distance among clumps or stands of trees that such treeless vegetation was grassland. These treeless (grassland) areas made up more of the total land area of the pasture than did that supporting trees. From that perspective the total or general vegetation of this pasture could be interpreted as that of a savanna. Conversely, it could be viewed as an overall woodland (as a combination of closed-canopy stands of oak and open grassland).

Freestone County, Texas. October; grain-ripe phenological stage in bahiagrass and Vaseygrass. No units of native vegetation were applicable for this naturalized woodland pasture. East Central Texas Plains- Floodplains and Low Terraces Ecoregion, 33f (Griffith et al., 2004).

150. Sward of Pineywoods naturalized pasture- Herbaceous layer of a naturalized woodland or savanna pasture on the western edge of Texas Pineywoods. Bahiagrass and Vaseygrass were the two major herbaceous species with bahiagrass almost always dominant or co-dominant with Vaseygrass which was, communitywide, the associate herb. Other grasses included the introduced species of common bermudagrass and Johnsongrass along with native grasses. These latter included perennial species of purpletop, broomsedge bluestem, beaked panicgrass, thinseed paspalum, and brownseed paspalum (in that approximate order) and the annual, Texas millet. There was some cover of caric sedges and umbrella or flat sedges. Forbs in this naturalized vegetation were limited to hogwort or wooly croton which grew in patches, none of which were in the sward presented here.

151. Bahiagrass (Paspalum notatum) on naturalized Pineywoods range- Turf and sexual shoots of bahiagrass (first slide) and bahiagrass panicle (second slide), the dominant herbaceous species on naturalized pasture on water oak-southern red oak woodland or savanna at western perimeter of Texas Pineywoods. The "dead giveaway" characteristic of the bahiagrass panicle is the physical arrangement in which the two spicate branches usually arise at a common point on the central axis of this inflorescence and then frequently curving symmetrically downward (as in the second of these slides). Sometimes one panicle branch arises slightly higher on the floral axis (ie. one branch is a "little higher up on the seed stalk" than the other branch). Bahiagrass is the only Psapalum species in the Texas Pineywoods that regularly has this point of common origin for main branches in its panicle (Gould, 1975, p. 501).

Freestone County, Texas. October; grain-ripe phenological stage.

152. Spikelets of the naturalized dominant- Ripened caryopses on branches of a panicle of bahiagrass, the dominant herbaceous species on naturalized water oak-southern red oak woodland or savanna range at western extremity of Pineywoods in east Texas.

Freestone County, Texas. October; grain-ripe to grain-shedding phenological stages.

153. Spikelets of the natralized associate- Inflorescence of Vaseygrass, an agronomic perennial grass that was the associate herbaceous species on water oak-southern red oak woodland or savanna having an understorey dominated by naturalized domestic grasses at western perimeter of Pineywoods in east Texas. Panicle and flag leaf were shown in first photograph while the second slide featured ripening spikelets.

The Paspalum inflorescence type has traditionally been regarded as a panicle. Contemporary terminology described the major units of this panicle as "unilateral spicate branches" (Gould, 1975, p. 500). These branches in P. urvillei are ascending or erect and closely spaced among themselves and arising off both sides of the central axis (or rachis) of the panicle.

Freestone County, Texas. October; hard-dough to grain-shatter stages of phenology (and all on the same branch of a panicle).

154. Texas millet or Texas panicgrass (Panicum texanum), a native, warm-season annual grass on naturalized (and, mostly, perennial) pasture- Local stand of Texas millet growing in the herbaceous understorey of water oak-southern red oak woodland or savannal range. The plants representing this species were growing on the example of naturalized Pineywoods grazing land shown and described above. Texas millet was not a major species in this pasture vegetation, but it was important and even dominant on some local sites (microhabitats).

Freestone County, Texas. October; soft-dough stage of phenology.

155. A native, warm-season annual and its sexual shoots- A single plant of Texas millet (first photograph) and two sexual shoots of this species (second photograph). This small to mid-size panicgrass is widespread in North America ranging from the Atlantic Coast (Massachusetts) to Pacific Slope (California). Texas millet is often a weedy or ruderal species, even in areas where it is a native. More specifically, it tends to be a pioneer (at least early seral) species on disturbed sites. The specimen presented here was growing in the West Cross Timbers on land subjected to repeated heavy (close) mowing.

Erath County, Texas. October; anthesis.

156. Spikelets of a native warm-season annual grass- Spikelets on contracted panicle of Texas millet. Different spikelets were in various stages of flowering and fruit formation ranging from anthesis to grain in milk-stage. Examples were from plants growing in the West Cross Timbers on overmowed land. This annual species was well-represented in a Texas Pineywoods woodland dominated by water and southern red oak in the tree layer and bahiagrass and Vaseygrass in the herbaceous layer. Texas millet added two floristic components to this understorey: 1) annual life cycle and 2) native species. Other native grasses and grasslike plants in this herbaceous zone of vegetation were perennials.

The inflorescence of Texas millet is made up of erect or ascending branches that lie closely to and spread slightly away from the central axis (or rachis) resulting in a "tight" panicle yet with branching conspicuous enough that it is not a contracted panicle.

Erath County, Texas. October.

157. A native annual forb on naturalized (perennial) Pineywoods range- Hogwort or wooly croton (Croton capitatus) on water oak and southern red oak-dominated woodland with an understorey composed largely of naturalized bahiagrass and Vaseygrass. Hogwort was the most common forb in the herbaceous layer(s) of this highly man-modified vegetation.

Freestone County, Texas. October.

158. Wooly croton or hogwort- Shoot apex of Croton capitatus showing fruit which is interpreted as a capsule composed of two to four carpels each of which contains one seed (Fernald, 1950, p. 959). All Croton species are monecious. The apt adjective "wooly" was clearly evident in this photograph.

Freestone County, Texas. October.

159. Water oak (Quercus nigra)- Leader (first photograph) and detail of leaves (second photograph) of water oak growing on a naturalized woodland or savanna pasture on the western margin of the Texas Pineywoods. Southern red oak was the co-cominant woody species on this highly man-modified grazing land. Bahiagrass and Vaseygrass dominated most of the herbaceous layer of this naturalized range. There was limited regeneration of oaks on this frequently close mowed (overmowed) pasture with tree reproduction limited to short seedlings or saplings growing next to trunks of established trees where they were protected from a tractor-mounted shredder.

Freestone County, Texas. October.

160. Southern red oak (Quercus falcata)- Leaves of the co-dominant tree on a naturalized (anthropogenic) woodland or savannah pasture on western edge of Texas Pineywoods. Bahiagrass and Vaseygrass were understorey dominants over most of this naturalized range. Water oak was co-dominant with southern red oak in the tree layer(s). Regeneration of Quercus species was limited to seedlings that had not grown to mowing height or where growing close enough to trunks of larger trees that they could not be reached by the tractor-mounted rotary mower.

Freestone County, Texas. October.

Disclimax Arizona Annual Grassland

161. Arizona annual grassland- Disturbance climax annual grassland of red brome, foxtail brome, or foxtail chess (Bromus rubens), dominant, and wild oats (Avena fatua), associate species, on a degraded semidesert grassland formerly dominated by black grama (Bouteloua eriopoda) with blue grama (B. gracilis) and galleta (Hilaria jamesii) the main associates along with some cover of sideoats grama (B. curtipendula). Potential natural (climax) shrub species of major importance were not known, but catclaw, catclaw mimosa, or wait-a-minute bush (Mimosa biuncifera) was about the only woody species on what was almost certainly a grazing disclimax (disturbance climax) atrtributable to overgrazing by livestock at some point in recent history of this range plant community). Another likely contributory factor was improper use of fire. This was more likely to have been fire exclusion rather than improper intensity, frequency, season, etc. of wild fire and/or anthropogenic fires. A third factor that contributed to development of this disclamax "Arizona annual grassland" was road-building activity. This range was conterminous with an interstate highway,and there was clear physical evidence that heavy equipment and eath-moving operations had adverse impacts on this arid grassland landscape.

This range was still being grazed by cattle as shown by presence of "cow brutes" in both of these wide-angle shots. Even a "quick look" at this range revealed that current degree of use of plants of these two annual grasses was relatively light (see slide and caption "two doors down"). Stocking rate or grazing intensity as measured or estimated by degree of use showed conclusively that current grazing was not (had not been) excessive, that is had not exceeded proper degree of use in the most recent past (the last or preceding) grass-growing season. Overuse had not taken place during the last cool-season. Based on plant residue (quantity of mulch or grass herbage, stubble height, etc.) grass utilization had been light. At this rate of defoliation (intensity or degree of use), by beef cattle in this instance, range degradation would not have occurred. Degree of use was light enough that even the extremely grazing-sensitive black grama would not have been eliminated.

By "reading range sign", however, it was obvious that past degree of use--at some point in time of white man's grazing use (history of livestock use) of this range-- overuse over a long time span had resulted in changes in the range plant community (ie. overgrazing by livestock had occurred). Such long-term overuse (= overgrazing) combined with 1) human introduction of Eurasian grass species that evolved under heavy grazing use, 2) likely changes in fire regime (fire exclusion, fire at seasons/stages of plant phenology injurious to native perennial grassess, etc.), and 3) road construction (this range was adjacent to an interstate highway with signs of damage by heavy, earth-moving equipment) resulted in replacement of the climax grama-galleta semidesert grassland by/with Eurasian annual grasses resulting in creation of "Arizona annual grassland".

This unintentional or de facto man-made type conversion was dubbed "Arizona annual grassland" by the author who had past research experience with California annual grassland. In both such cases, type conversions followed the same pattern of human disturbance that resulted in a disturbance climax, (Clements, 1936, p. 265; Weaver and Clements, 1938, ps, 86-88). This disclimax "Arizona annual grassland" is, in this rangeman's judgment, permanent in human time scale just like it is for the California annual grassland type. The current grazing use of this annual grassland range was proper. It was wise use at rates on the light side of sustained yield. Nothing would be gained by failure to use the feed resource. In fact, herbivory by domestic cattle might facilitate ecosystem function by increasing rate of herbage decomposition or planting seed of now-permanent, naturalized, Eurasian grasses.

Yavapai County, Arizona. Mid-July; dead shoots, live caryopses (senescence, mature fruit) at end of annual life cycle. Description of this range vegetation was not to be found in the literature (except for California and the Middle East). Cinder Hills range site.

162. One native (and well-armed) species remained- Catclaw, catclaw mimosa, or wait-a-minute bush (Mimosa biuncifera) was about the only native plant species that remained on a grazing disturbance climax annual grassland of red brome (dominant) and wild oats (associate) on what had been climax black grama-blue grama-galleta semidesert grassland. For all practical purposes, catclaw was the only native plant species remaining on this deteriorated range that was now in a state of disturbance climax. The anthropogenic successional state of disclimax was explained in the preceding caption. In short (no pun intended), the native warm-season, eragrostoid climax grasses had been "grazed out" (probably under unregulated open-range grazing on what would have been Public Domain land) leaving range resources of soil (what remained after accelerated [soil] erosion), limited precipitation of the Arid Zone, light, carbon dioxide, etc. available to Eurasian annual grasses that had been introduced by European man after these cool-season (festucoid) grasses had evolved for millenia under heavy livestock grazing in the Mediterranean Region. The only native range plant species that fared "ok" under such explotiative, improper grazing land management was the prickle-equipped, low-palatability catclaw.

Wait-a-minute is viciously armed with talon-like (catclaw-shaped) prickles that occur both singularly and in pairs and that ward off most would-be browsers. Mimosa biuncifera is widely distributed in North America being a component of Great plains grasslands and extgending westward through the Chihuahuan and Sonoran Deserts, and, even some, in the Mojave Desert. Catclaw mimosa is probably most abundant in the semidesert grassland and adjoining shrubland range types like the "soft chaparral" in mountain fooothills.

Good--though basically brief--discussions of M. biuncifera included Dayton (1931, p. 79), Vines (1960, ps. 507-508), and Benson and Darrow (1981, ps. 232-233). Apparently the prickles of this species are a deterrent to browsing animals (other than for legumes which are quite palatable even to livestock), but catclaw was rated as Fair in feed value during winter and drought by Dayton (1931, p, 79). Benson and Darrow (1981, p. 233) concluded that catclaw mimosa was readily eaten by whitetailed deer but provided "little sustenance to livestock".

Mimosa species are legumes of the mimosa subfamily (Mimosoideae) which generally do not fix nitrogen. It is the papilionaceous leguimes (subfamily, Papilionoideae) that are the nodulated legumes, those that enter into mutualistic interrelationships with nitrogen-fixing bacteria (primarily Rhizobiium species). As shown in the preceding paragraph there is not a lot of information or basic knowledge concerning Mimosa biuncifera.

Presence of catclaw mimosa on this disturbance climax "Arizona annual grassland" created a savanna or, at least, a savannah-like physiogonomy to this disclimax range plant community.

Yavapai County, Arizona. Mid-July; prebloom phenological stage of Mimosa biuncifera; dead shoots, live caryopses (senescence, mature fruit) of annual grasses. Description of this range vegetation was not to be found in the literature (except for California and the Middle East). Cinder Hills range site.

163. Mediterranean disclimax- Sward of red or foxtail brome and wild oat(s) on a once-upon-a-time black grama-blue grama-galleta semidesert grassland that through human action (introduction of alien species, overgrazing, underburning, road construction, etc.) was degraded to an annual grassland range with red or foxtail brome as dominant and wild oats as associate species.

This "photoplot" of grazed annual grass herbage was on the range presented in the two immediately preceding slide-caption sets. Degree of use had been proper for this grassland range feedstuff from standpoint of both 1) grass plants (current and the next generation or annual life cycle) and 2) nutrition, productivity, and health of cattle grazing this range. Students should note that there were some remaining (ie. ungrazed) sexual shoots with spikelet-filled panicles of the two annual festucoid grasses. There was adequate grain crop remaining to insure next season's crop of cool-season annual grasses, a crop of a disturbance climax that is "here for the duration" on human time scale.

It was explained above that overgrazing by livestock, altered fire regimes by whiteman, and road-building activities had all contributed to degradation of the former climax black grama-blue grama-galleta semidesert grassland with its conversion to a disturbance climax (disclimax) of red brome-wild oats annual grassland. This anthropogenic grassland was the result of an unintensional type conversion: climax desert plains grassland (Clements, 1920, ps.144-149; Weaver and Clements, 1938, ps. 525-526) converted to "Arizona annual grassland", the ecological/successional equivalent of the long-recognized (and known to be permanent) disclimax of California annual grassland. Make the best of it. That is the essence of sound (proper) range management. Sadly, the black grama "ain't coming back", at least not on a time scale relevent to human beings.

Red brome has become the southern or hot desert ecological equivalent of cheatgrass (Bromus tectorum) in the Intermountain Region and Japanese chess (B. japonicus) in the central North Amrican grasslands. Invasion by red brome, especially the degree or extent of invasion, was an invasive development that was apparently more recent than that of the other naturalized, Eurasian, annual Bromus species. For example, Humphrey 1960, p. 31) explained that while red brome was "[p]articularily common on ranges where the original perennial grass cover has deteriorated" the species "appears to stilll be spreading in Arizona". Nontheless, red brome was recognized as a major, naturalized grass by Gould (1950, p. 50) who described this weedy Eurasian annual as being "abundant over large areas in western United States, especially on overgrazed rangelands".

Gould (1950, ps. 50-51) described the herbage of red brome as being "[o]f some forage value, especially for sheep, but with an extremely short period of palatability".

Again, "it ain't much", but sound resource management is to make the best use of what is,and will continue to be, there indefinitely. If we could turn back the ecological clock... But we cannot. The Moving Finger of Omar Khayyám has writ, and "...having writ, Moves on".

164. Red alien -Small group of several individual plants (first slide) and two individual plants (second and third slides) of red brome, foxtail brome, or foxtail chess (Bromus rubens) growing in the Arizona Trransition Zone or Central Highlands (between the Colorado Plateau and Basin and Range physiographic provinces). Red brome is another of the Eurasian annual Bromus species introduced (mostly accidentally) into North America during the early days of New Spain. At the present time the now-naturalized red borme appears to be increasing (density, frequency, cover, etc.) within its new home (its species range of naturalization) on North American ranges. Currently it seems that red brome is more-or-less the ecological equivalent in the arid zone of southwestern North America to cheatgrass or downy brome (Bromus tectorum) in the Intermountain Region and Japanese chess (B. japonicus) in the grasslands of the Central Lowlands and Great Plains.

Red or foxtail brome has become the dominant annual grass throughout much of the Mojave Desert and much of the Sonoran Desert (including the Lower Colorado Valley subdivision, commonly regarded as the "Colorado Desert"). Red brome appears to be increasing its general abundance within the Arizona Upland subdivision of the Sonoran Desert. Apparently this increase has been both 1) an expansion of red brome's biological (species) range expansion and 2) an increase abundance throughout former parts of its natrualized range as well as newly expanded portions of this range of naturalization. nhas been For instance, Shreve and Wiggins (1964, p. 241) did not list B. rubens in their group of Bromus species for the flora of the Sonoran Desert. Kearney and Peeples (1951, supplement 1960; p. 78) did list B. rubens as occurring in "roadsides and waste places" in part of Arizona. Likewise, Gould (1951, ps. 50-51) showed both B. rubens and B. madritensis as being reported from some of the desert counties of Arizona. In Arizona Range Grasses Humphrey (1960, p. 31-32) and, in the revised edition of this bulletin, Ruyle and Young (2003, ps. 42-43) observed that red brome appeared to be spreading in Arizona.

Red or foxtail brome appears to pose some threat--whether this is minor or considerable remains to be determined--to native range communities, both from plant and animal perspectives. This has been most discussed for desert range ecosystems. Students can find more such discussion of red brome on desert ranges in the chapter, Sonorad Desert.

Montezuma Well unit, Montezuma Castle National Monument, Yavapai County, Arizona (first slide); Yavapai County, Arizona (second and third slide); Mid-July, standing dead at grain-ripe stage of phenology.

165. Resource allocation for aridity- Shoots (first slide) and panicles (second and third slides) of red brome in the Transition Zone or Central Highlands of Arizona. This naturalized, Eurasian annual grass produces amazing quantities of grain having as many as eleven florets per spikelet (Hitchcock and Chase (1950, p. 54). Wu and Jain (1979) found an average of 76 seeds per plant of red brome. Bromus rubens is closely related to B. tectorum, B. mollis, and B. rigidus all of which are naturalized Eurasian annuals that Hitchcock and Chase (1950, p. 33) included in the section Eubromus. B. rubens is the shortest in stature of these other Eubromus species.

B. rubens is closely related to B. madritensis but these taxa were kept as two species in Barkworth et al. (2007, ps. 226-227).

Genotypic and phenotypic aspects of red brome were reported by Wu and Jain (1978) and population regulation (including life cycle and competition) of red brome was described by Wu and Jain (1979) while natural history and ecological aspects of red brome was summarized by (Newman 2002).

Yavapai County, Arizona (first and third slide); Montezuma Well unit, Montezuma Castle National Monument, Yavapai County, Arizona (second slide). Mid-July, standing dead at grain-ripe stage of phenology.

Time-tested references: In addition to Arizona Range Grasses (Humphrey, 1960; revised by Ruyle and Young, 2003), which dealt with desert grasses from a range management perspective, another outstanding source for general and, especially, taxonomic treatment of aridlands Gramineae was Grasses of Southwestern United States (Gould, 1951).

Summary of grass species photographed in the Arizona Upland vegetational subdivision: based on presence of eragrostoid and panicoid (both annual and perennial) grasses and review of the classical literature, it seemed a valid conclusion that there is an herbaceous layer in the Sonoran Desert vegetation described as rangeland cover type SRM 507 (Palo Verde-Cactus). Native perennial and annual grasses, along with naturalized Eurasian annual grasses, are frequently or occasionally part of the Sonoran Desert vegetation. Some of these same grass species also occur on range dominance types that are conterminous with (or in close proximity to) Arizona Upland Sonoran Desertscrub (eg. semidesert grassland and Chihuahuan Desert). These grasses-- both native and naturalized Mediterranean species-- are part of the potential natural vegetation of all these range cover types as well as that of the vegetation units mapped and described by Kuchler (1966, 1971) and Brown (1994).

Recent descriptions of the natural vegetation of the Arizona Upland subdivision have in effect ignored the herbaceous layer. Descriptions dealt almost exclusively with woody plants (including succulents) that dominate the upper or open canopy, middle, and lower layers (ie. upperstorey and mid-stories of trees and shrubs, including cactus and succulents). There has been either very little or no description and discussion of the herbaceous layer which this author concluded to be part of the structure of climax or potential natural vegetation. For example, the Society for Range Management description of the Palo Verde-Cactus cover type made mention only of herbaceous plants that were annual grasses and forbs (most of these were naturalized Eurasian species). These naturalized Mediterranean species should be interpreted as part of the now-potential (or climax) vegetation. They may have impacts both positive (increase forage production or diversity of animal diets, increase biological diversity) and negative (compete with natives for limited resources, increase incidence of unnatural fire regimes). Eurasian species are not, however, the only grasses and forbs in the Arizona Upland vegetation. In fact, they are not necessarily the dominant, most common, or defining herbs. Descriptions of herbaceous species in Arizona Upland range vegetation should not stop with naturalized exotics.

The classic and seminal literature on Sonoran Desert vegetation, most notably that written by Forrest Shreve, gave more detailed description of the herbaceous layer and species comprising it-- both perennial and annual-- than recent works. This should be rectified. The herbaceous part of Sonoran Desert range, however limited (eg. sparse in several of the natural range communities in the Lower Colorado Valley and Arizona Upland), is the defining part from the perspective of the vegetation as range. This herbaceous understorey is the key part of Sonoran Desert vegetation from the standpoint of using these forage and resources by scientific range management.

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